A HANDB O OK OF T H E WOR L D’ S C ON I F E RS

Cupressus guadalupensis Drawing by Aljos Farjon

A HANDB O OK OF THE WORLD’ S C ONIFE RS

by

AL JO S FA R JON

Volum e I

BRILL
Leiden-BOSTON
2010
 This book is printed on acid-free paper.
Library of Congress Cataloging-in-Publication Data
The Library of Congress Cataloging-in-Publication Data is available from the Publisher.

front cover:
Abies fabri young seed cones

back cover:
top left: Dacrycarpus kinabaluensis seed cones
bottom right: Picea likiangensis young seed cones

isbn: 978 90 04 17718 5

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TABLE OF CONTENTS

Foreword 7

Preface 9
5
The conifers of the world, an introduction 13

The distribution and ecology of conifers 21

The economic importance of conifers 25

The conservation of conifer diversity 29

Synopsis of families and genera 33

Taxonomic treatment of families, with keys to families 43

Taxonomic treatment of genera and species 55

(in alphabetical order), with keys to sections, subsections and species

Glossary 1033

References 1053

Lists of illustrations 1065

Index to botanical names of conifers 1073

FOREWORD
With the Handbook of the World’s Conifers, its au- because it gives us the keys to designing effective
thor Aljos Farjon has given us a wonderful aid to conservation measures for each endangered species
learn almost all there is to know about this ancient and also defines the scope and limitations of where
lineage of woody plants. In true Handbook tradi- one could introduce a species successfully in tropi-
tion all 615 species of extant conifers are concisely cal or temperate horticulture. 7
described, with user-friendly identification keys to
families, genera and species and invaluable addi- With his lifelong dedication to the study of the tax-
tional information on uses, conservation status, and onomy of conifers at Utrecht, Oxford and Kew, and
much more. Unlike earlier books on conifers that his strong involvement in the conservation of many
are heavily skewed towards temperate conifers from red-listed conifer species throughout the world,
the northern hemisphere, the present Handbook Aljos Farjon is arguably the only plant scientist in
does equal justice to the 200 tropical species and the world to complete the colossal task of produc-
415 temperate ones; this attention to tropical species ing this comprehensive handbook single-handedly
makes the Handbook truly unique. and authoritatively. I hope and am confident that it
will give as much pleasure and satisfaction to a wide
In the very informative introduction the evolution, ­audience of tree enthusiasts, foresters, botanists,
classification, ecology, biology, economic uses, and ecologists and conservationists as it has given me.
conservation status of the conifers are discussed.
This handbook thus combines the virtues of a com- Pieter Baas
prehensive taxonomic monograph of the conifers, Emeritus Professor of Systematic Botany, Leiden
with that of a true vademecum on the morphologi- University
cal attributes, uses, and conservation of all species Former Director of the National Herbarium of the
of the order. The information on the ecology of each Netherlands
species in its natural habitat is particularly welcome,
PREFACE
Many books have been written about conifers dur- graphs on ecology, uses and conservation with all
ing the 19th and 20th centuries, the majority of these species accounts. This handbook does not describe
with emphasis on horticulture in Europe. Those that or illustrate conifers that are only known in cultiva-
are mostly compilations of species and their culti- tion. With some 15,000 conifer cultivars known, no
vars found to be grown in gardens and parks, with comprehensive coverage could have been achieved. 9
descriptions, illustrations and information about Instead, a separate volume now covers these,
their cultivation, are known as handbooks or manu- ­authored by Aris Auders in Latvia and Derek Spicer
als. A conifer handbook as traditionally compiled in England, entitled Conifers – A Comprehensive
is therefore a comprehensive guide to the conifers Guide to the Conifer World, published under the
that are grown or could be grown in cultivation in a auspices of the Royal Horticultural Society. It fol-
certain part of the world and includes, besides spe- lows the taxonomy in this Handbook of the World’s
cies and botanical varieties, cultivated varieties or Conifers for families, genera and species and these
forms (since 1953 universally known as cultivars). two handbooks are therefore complementary and
All such works, even the most comprehensive, em- can (and should) be used side-by-side.
phasise conifers adapted to a cool temperate climate The need for a comprehensive and modern hand-
as experienced in Europe. Several have made this book concentrating on the species of conifers is clear.
explicit in the title of the book, but even those that Awareness of the obligation to manage and protect
have not done this invariably treat ‘hardy’ conifers the biological resources of this planet and use these
more fully than other species, which are at best only resources sustainably has created a demand for ac-
mentioned briefly. The scope, content and presen- curate, up-to-date information. Conifers occur
tation are aimed at the horti- or silvicultural users. worldwide and are major or minor components of
Some of the better-known conifer handbooks were nearly all major forest and woodland types, as well
reviewed in Chapter 1 of my Monograph of Cupres- as many timber plantations and numerous gardens
saceae and Sciadopitys (Farjon, 2005a). Most coni- and parks. They are of high economic value, but
fer handbooks are now out-of-print, but some new also play major roles in the ecology of many natu-
books have recently been added to the bibliography ral and semi-natural ecosystems. They are an inte-
of conifers, while others are being compiled. The gral part of many landscaped estates and parks from
emphasis on the temperate conifers in cultivation the subtropics of Australia and New Zealand to the
has remained strong in these works, as far as I know. temperate regions of Europe and North America.
The present Handbook is very different from This handbook of conifers has been compiled to ad-
these predecessors. There is no particular emphasis dress this need for relevant information and should
on conifers that can be grown in Europe. Of the 615 therefore be useful to botanists, conservationists,
species known in the world at present, about 200 oc- dendrologists, ecologists, foresters, horticulturists,
cur in the tropics. They receive here equal cover with land resource managers and, I should add, all those
the species that grow in temperate climates. In this with an interest in the trees and forests of the world.
respect, the present book more closely resembles a Knowledge is not only useful, it is a pleasure that
taxonomic monograph, as it includes all the current- enhances the quality of life.
ly known and accepted taxa, from the ranks of fam- The introductory chapters have deliberately been
ily down to variety and forma. However, it is not a kept brief, often merely touching on interesting as-
monograph purely for taxonomists. Its content aims pects of conifer biology, evolution and taxonomy, as
at a much wider audience. To achieve this, strictly well as ecology, economics and conservation. For
taxonomic information has been limited to what is a recently published full introduction I refer to my
more or less essential, while other information has book A Natural History of Conifers (Farjon, 2008),
been included that may not even have been consid- where all these issues are treated in much more de-
ered for a taxonomic monograph. There are para- tail. After these short chapters, the book moves to
 the families, genera and species and becomes the nyms) or begin on a new line. Where relevant, the
handbook referred to above. For the general user it (aberrant) status of synonyms under the Botanical
may be useful to outline and explain the conventions Code (ICBN) is given after the citation of the place
and format followed throughout for all genera and of publication. Types (i.e. herbarium specimens des-
species. To identify a completely unknown conifer, ignated to fix the application of a botanical name)
one should start with the key to families on p. 43; are only given here for accepted names, including
then the key to genera under the family determined, their synonyms if based on the same type. Types of
and the key to species under the genus arrived at. genera are cited as names of species. The opportu-
Genera with numbers of species greater than 15–20 nity to designate types where it was appropriate and
have been divided, formally or informally, and keys possible has been taken; but in several cases typifica-
10 to these groups are first to be consulted. Under spe- tion is complicated and then it has been left to future
cies may follow subspecies and/or varieties; these research. This is what is meant with the phrase “type
are usually not included in keys. Not more than six not designated” under several species. Type speci-
infraspecific taxa are accepted under any one spe- mens are deposited in institutional herbaria, abbre-
cies in this book, and their descriptions are brief viated by an acronym and these can be found at:
and diagnostic. These descriptions should primarily www.sciweb.nybg.org/science2/IndexHerbariorum.asp
be compared with the relevant species description Under species, subspecies, varieties and forms
and the character states mentioned therein, and may be recognized. While these ranks are hierarchi-
less with each other. No attempt has been made to cal in that order and species have been subdivided in
create varieties within subspecies and, unless such that way, I am no advocate of this. A single rank un-
ranking has been established by other authors and der species would suffice in botany as well as it does
is followed here, the two ranks are treated as equal. in zoology. Leaving the rare use of forms aside as a
In the taxonomic literature, the use of either subspe- special case, I am agnostic as to whether this should
cies or variety appears to derive more often from be the rank of subspecies or variety. In botanical
tradition than from explicit taxonomic concepts. taxonomy, we have an unavoidable historical legacy:
I believe that botany would be well advised to follow both ranks have been used in the literature, often
zoological practice and use only a single infraspe- for the same taxon. In some countries one rank was
cific rank in future. After using the keys, reading the preferred over the other. In this Handbook, there-
description of a species is recommended as a check fore, I have used what ranks were available in validly
to find out if the determination is correct. Also, as it published names under species where infraspecific
is almost impossible to construct keys that use any taxa merited recognition; under a few species I have
and every feature of a specimen in hand, the reading accepted two ranks if these appear to be widely used.
of the description helps with the identification. An Unfortunately, the same notion of one rank under
extensive glossary is provided to help explain much species has induced the publication of numerous
of the terminology used in this handbook. new combinations, some have changed subspecies
Botanical names are given in full as in a taxo- to variety and others vice-versa depending on prefer-
nomic monograph, i.e. with (abbreviated) author ences. I have not seen fit to increase the number of
name(s), abbreviated reference to the place of first pages by including all of these mostly superfluous
publication, synonymy and type citation. The ac- name changes, nor by adding new combinations if
cepted names are followed by synonyms if these none existed in the desired rank. If infraspecific taxa
­apply. Among synonyms, basionyms are always are recognized, the species name in the inclusive
cited as types refer to these; other synonyms are sense is given first (with ­authority and place of publi-
cited selectively. In most cases, synonyms no longer cation) and the description is similarly inclusive, i.e.
in use as accepted names have been omitted. Most it encompasses all subspecies and/or varieties. Syno-
recently published names have been included, but nyms and types are given with the latter and the short
not all, for instance new names resulting from a descriptions given there are exclusive. Botanical de-
mere change of rank, such as variety to subspecies. scriptions follow the names and synonyms of taxa
When several synonyms are cited, homotypic syno- under the heading Description and other informa-
nyms are grouped together in one continuous line; tion is likewise given under its specific head for clar-
hetero­typic synonyms are separated by a blank line ity and ease of reference. The descriptions of species
from the accepted name (and its homotypic syno- follow the conventional sequence for ­descriptions of
trees; i.e. from size and habit to trunk, bark, branch- www.catalogueoflife.org/search.php
es, foliage, leaf details, male and female reproductive Ecology describes the habitat of a species and its
structures and ending with seeds. Despite the rela- altitudinal range (in meters above sea level: m a.s.l.)
tive detail in some descriptions, it should therefore is given here. For many species, especially in the
not be difficult to find particular parts and I have tropics, their habitat remains poorly documented,
therefore not marked these with a different font or while much has been written on many species in
type. The botanical names of species are followed by the temperate northern hemisphere. Some informa-
the etymology of the accepted name and by vernac- tion could be assembled from notes on herbarium
ular names. Etymology is also explained for genus specimen labels and has been recorded in the coni-
names but not for subspecies and varieties and only fer specimen database available at dps.plants.ox.ac.
occasionally vernacular names are mentioned with uk/bol//home/default.aspx under “Online Groups”. 11
the latter two. The vernacular names do not provide As indicated above, a balanced approach has meant
a comprehensive listing, especially not for species that not all that is known can be mentioned here
with a very large natural distribution, covering an and for such species the information on ecology
area in which many languages may be spoken. Ver- (and other subjects) is indicative rather than com-
nacular names may not be specific; if referring to the prehensive. Conservation is usually concerned with
genus rather than the species and if more than one species survival issues as defined by the World Con-
species occur in the area, such names are not listed. servation Union’s (IUCN) Species Survival Com-
Some published vernacular names seem contrived; if mission (SSC). Information was gathered and the
these are not actually in use locally or regionally, they conservation status of taxa (species to varieties) has
are not cited here. Taxonomic notes may follow the been assessed by members of a Specialist Group
descriptions; they are usually comments on different (SG) resorting under the SSC. In the case of conifers
taxonomic treatments or notes on typification of the this is the Conifer Specialist Group (CSG) which is
species or mention similarities of taxa. In the con- currently chaired by this author. Almost all conifers
cise format necessary in this Handbook, comments have been assessed under IUCN Red List criteria,
and discussions are limited to the essential and are the resulting categories of threat are given under
kept brief. Not every name placed in synonymy can the heading IUCN. For the abbreviated categories
be discussed here; the reader can rest assured that of threat see the introductory chapter Conservation;
due scientific consideration has been given, some of for definitions and decoding of the criteria used visit
which has been published elsewhere. Distribution www.iucnredlist.org/static/categories_criteria. The
of taxa is given in two formats and these are often listings given here are as in the 2008 IUCN Red List
complementary. The first is descriptive, mentioning of Threatened Species™ [www.iucnredlist.org] in-
regions as well as countries, provinces, areas and lo- cluding the criteria of either version 2.3 (1994) or 3.1
calities, as appropriate. The second format uses the (2001) as applied. Uses mentions commercial as well
TDWG codes (Brummitt, 2001). These codes pro- as other human uses of species. The two main cat-
vide geographical information at three levels defined egories are wood (timber) and its applications and
by geographically and politically delimited areas, e.g. horticulture, but other uses, including ‘traditional’,
42 = Malesia (a geographical region) BOR = Borneo have been mentioned when known. As with ecology,
(an island) and SB = Sabah (a politically delimited much more is known about uses of species in the
area): 42 BOR-SB means therefore that the species temperate zones than in the tropics. Often, timber
occurs (also) in Sabah. The meaning of these codes is traded under a ‘generic’ name which may even in-
can be found at www.catalogueoflife.org/search.php. clude different genera, as in the family Podocarpa­
For species with large ranges the TDWG codes pro- ceae, which makes distinctions of use among species
vide more detailed information than the descriptive extremely difficult. Horticultural use is also uneven-
statement, but for species with very limited distribu- ly distributed along similar lines of geographical de-
tion the latter is often more precise because TDWG marcation, but uses in the subtropics and tropics are
codes often do not go down to such detail. When picking up and therefore mentioned when known.
entered in a database, the TDWG codes linked to This book does not include cultivars but if such exist
names can provide easy and quick listings of species under a species, that information is given. Planta-
per country, state or province. This information is tion in a forestry context is usually mentioned with
available for conifers at:
 the uses of timber; for several species plantations funds. This was generously undertaken by the Lin-
have become the main source of that commodity. nean Society of London and I am grateful for the
The glossary in this Handbook is the most inclu- trust the Society has placed in one of their Fel-
sive I have yet compiled. I have made use of several lows to fulfil the commitments thereby generated.
published in my earlier works, but added numer- The Treasurer of the Society, Prof. Grenville Lucas,
ous entries specifically for this book. In particular, guided me through this whole process in his capable
I have attempted to explain terms used by plant and incomparable manner and I am most indebted
taxonomists, because this book is meant to be used to him for this generous support. Most importantly,
and consulted by many others, amateur as well as my gratitude goes to all the generous donors to this
professional. I hope it will be found useful, but I project, who together have made this book possible.
12 am aware of the difficulty of finding a balance be- I have permission to name them here and shall do
tween explaining commonly understood terms and so in alphabetical order. They are the Arboricultural
thereby underestimating my readers and using un- Association (UK), Mr. Lawrence Banks (UK), The
explained terms for which one might need to con- Dendrology Charitable Company and the Interna-
sult a large dictionary. Finally, I have only included tional Dendrology Society, Dr. Barry Denyer-Green
botanical names of conifers in the index. Numer- (UK), the Lord Devonport Charitable Trust (UK),
ous other species names occur in the texts on the Mrs. Francine von Finck (Germany), Mrs. Arabella
ecology of conifer species, but giving information Killander (UK), the Samuel Storey Family Charita-
about these other species is obviously beyond the ble Trust (UK) and the Stanley Smith Horticultural
scope of this Handbook. Vernacular names, while Trust (UK). This Handbook is the synthetic product
cited under many species, are incomplete (espe- of 25 years of research into conifers by its author. To
cially in the tropics), use several languages besides give detailed acknowledgement to all persons and
or in lieu of English, and are often ambiguous. The organisations who have assisted me in that research
reader is therefore encouraged to use (and learn) the would be next to impossible. Research is often a col-
botanical names of species. Both in the library and laborative effort, even if the resulting published out-
on the Internet, nearly all information is available put appears to have a single author. My thanks are
under these identifiers. Because the information in therefore here expressed to all I have been in touch
this book is arranged by species, finding the species with on conifers from time to time, on field trips
(they are arranged in alphabetical order as well as around the world as well as at my and their various
indexed) is all that is needed to obtain all the infor- home institutes during those years. Dr. Christopher
mation about them given in this Handbook. N. Page reviewed the texts on Podocarpaceae and
The compilation and writing of the Handbook the smaller families and his supportive comments
of the World’s Conifers has been a task which were much appreciated. Michael Frankis comment-
started some time after my official retirement from ed on common names and corrected errors therein.
the Royal Botanic Gardens, Kew in 2006. Certain The entire text was copy-edited by Dr. Hans Kruijer,
conditions had to be met before I could seriously whose helpful corrections much improved consist-
embark on this project, because it required access ency and clarity of presentation. Prof. Pieter Baas
to resources, to enable me to devote full work- wrote a thoughtful foreword, for which I am most
ing time to it. I was very fortunate that the Royal grateful. Finally, coming closer to the end product
Botanic Gardens appointed me an Honorary Re- now before you, I wish to thank the publisher, Brill
search Associate after my retirement, to be based and their helpful staff (especially Michiel Thijssen,
in the Herbarium where I had worked before (now Sabine Steenbeek and Frits Fritschy) for their pro-
the Herbarium, Library, Arts & Archives Depart- fessional work and helpful patience with the author.
ment). It was there that much of the work on this They have turned a huge and complicated manu-
book was done and I am grateful to RBG Kew for script into a beautiful book.
this invaluable support. Funds had to be raised for
this project, and when money was being pledged, I Aljos Farjon, FLS
had to find a registered charity to administer these [Kew, 27 November 2009]
THE CONIFERS OF THE WORLD, AN INTRODUCTION
To most of us who live in the temperate zone of the But, such criteria of appeal to biologists aside, the
northern hemisphere, conifers are among the most sheer number and volume of conifer trees growing
familiar trees and indeed an everyday sight. Not in the temperate north makes them of prime eco-
only do they naturally occupy vast areas of north- nomic and ecological importance throughout the
ern lands such as Canada, Scandinavia and Russia, entire world. With an estimated value of $100 bil- 13
as well as many mountain regions further south, lion per annum, wood products stand foremost and
we have also extensively planted them in forestry conspicuous at the top of the economic shopping
while gardens and parks of even moderate size are list, but especially at more southern latitudes other
hardly conceivable without them. Far less widely products, e.g. resins and derivatives, and even seeds
known are the conifers of the southern hemisphere for food, are also economically important. And
and still fewer people will associate conifers with then, as already indicated, there is the significant
the tropics. True, neither of these major parts of contribution conifers have made to horticulture as
the world have extensive conifer forests similar to ornamental trees, a trade which has given rise to nu-
those in the north; conifers in these latitudes are merous cultivars some of which are among the most
more often than not a relatively minor component popular shrubs and trees planted almost the world
of forests, often confined to rather inaccessible ar- over in gardens and parks. I venture the statement
eas, or inconspicuously mixed with more numerous that, although being a relatively small group of spe-
angiosperm trees. But if we were to regard diversity cies of woody plants ‒ there are about as many spe-
rather than sheer quantity as a criterion, the north cies of oak (Quercus) and twice as many species of
would lose its primacy as the realm of conifers to wattle (Acacia) ‒ conifers far exceed any other group
more southern latitudes. In the present Handbook of woody plant species in economic and ecological
of the World’s Conifers I have recognized 615 species importance. We would have to look at the major
in 70 genera belonging to 8 families. Of the genera, food crop plants to find species that generate more
35 are restricted to the northern hemisphere, 25 oc- income than some of the pines and spruces. But, as
cur only south of the equator and 10 are found both with cereals and beans, only a tiny minority of spe-
north and south of that line. Since, with the excep- cies diversity within conifers is accountable for this
tion of Podocarpus, which is largely tropical, most prominence today, while the majority shares in the
large genera occur in the northern hemisphere, that fame only by merit of kinship. Having said this, the
half of the globe (with by far the greater land mass) question may be asked, what constitutes this ‘kin-
has more species than the southern hemisphere. But ship’ in conifers? In other words, what are conifers?
in the northern hemisphere, diversity of species, It is not as straightforward a question to answer as
especially in the larger genera, increases markedly it may seem.
towards the equator, with for instance only 9 spe- In the early days of systematic botany, the ‘coni-
cies of Pinus in Canada and 43 in Mexico. More or fers’ were often thought of as a natural family of
less limited regions with more than 40 species or 50 plants, in a similar fashion as botanists referred to
taxa (species and lower ranks) defined as centres of e.g. Rosaceae or Compositae and, indeed, the name
diversity for conifers (Farjon & Page, 1999) are now Coniferae prevailed well into the early 20th century.
eight in number, seven are situated on or north of Are conifers a family of plants, or at a rank higher,
the equator and one south of it, but all except Japan an order Coniferales? Modern classifications classify
are at latitudes below 40° and two, Borneo and New groups of organisms in monophyletic groups, i.e.
Caledonia, are entirely in the tropics. That last island organisms or groups of organisms that in evolution-
is the most diverse and remarkable conifer centre of ary history share a common ancestor (usually an an-
all; covering an area the size of Wales, it has 43 spe- cestral species). Monophyletic groups may include
cies and all are endemic to the island. more recently formed monophyletic groups and be-
long to other monophyletic groups of more ­distant
 pasts. We must discuss the term ‘conifer’ in this con- a­ ngiosperms form a natural group. The term gym-
text of monophyly to answer these questions. No nosperm, on the other hand, does not specify a
such discussion makes any sense without looking at ‘natural group’ by any modern criterion. Conifers
the origin of conifers as (imperfectly) revealed in the are characterized by ‘naked’ ovules, for sure, and are
fossil record. The fossil story is a complicated tale thus gymnosperms, but they share this one feature
which can only be very summarily related here, but of early seed plants with other groups of primitive
I shall nevertheless try to answer these questions; for seed plants, e.g. Ginkgo and cycads. Taxa are de-
more on this topic, see my book A natural history fined by evolutionary novelties. It is, however, often
of conifers (Farjon, 2008). The word ‘conifer’, mean- difficult to determine which characters represent
ing ‘cone-bearer’, suggests that plants bearing ‘cones’ such evolutionary novelties. Taxa, therefore, need
14 could be called conifers. Cones are reproductive to be defined by diverse and structurally unrelated
organs consisting of fertile scales (simple cones) or characters to increase the likelihood of represent-
sterile and fertile scales (compound cones) that are ing ‘natural groups’ and for conifers several of such
spirally arranged at a central axis; the fertile scales characters can be identified. Farjon (2008) charac-
of a cone contain either pollen sacs or ovules. How- terized conifers as “shrubs or trees with secondary
ever, although the majority of conifers are charac- wood build of tracheids with large bordered pits in
terized by having cones, it is problematic to define their walls and narrow rays. The leaves are simple
the conifers as plants bearing cones. Even if the term and single or parallel veined. Resin is produced in
‘cone’ would be restricted to the ovulate organ of its the wood or in the leaves and is conducted through
description, a definition of conifers based on that resin canals. The reproductive organs are separated
organ is problematic, because we will then find that, into male and female, with male cones (pollen cones)
among the gymnosperms, pines are considered con- simple and female cones (seed cones) compound or
ifers but cycads are not. Conversely, Cephalo­taxus, reduced. Conifers have only one copy of a large in-
with a mature ovulate organ reduced to a single verted repeat in the chloroplast DNA, whereas all
seed is still a conifer, but Ginkgo, seemingly similar other plants studied so far have two copies.” If the
in that respect, is not (despite its inclusion in many DNA observation holds true, it may well turn out to
books on conifers). The key to the difference here is be the single character state that is unique to coni-
partly revealed in the ontogeny of both taxa, but it fers. But the group is exclusively circumscribed by
ultimately has to do with phylogeny, i.e. how these this plus the other characters, each not unique, but
taxa are related in an evolutionary sense. in combination only present in conifers. Such char-
acters are sometimes called ‘traits’. The seeds of cer-
The term ‘gymnosperm’ has fallen, and we need to tain coniferous taxa occurring in the Carboniferous,
explain what is meant by it first. In the course of the when the earliest seed-bearing plants became appar-
early evolution of land plants, adaptations to water ent, tended to be borne on appendages axillary to
stress ‒ i.e. dry conditions ‒ gradually led to the ori- leaves. Reduction of these appendages in size and/
gin of the ‘seed habit’, with the evolution of ovules or number, planation, movement of ovules towards
with an envelope (integument) to protect the vulner- the base of the appendage and transformation of the
able gametophyte phase from desiccation, coupled subtending leaf to a bract as well as reduction of the
with devices to catch pollen in a suitable micro-­ fertile shoot to a determinate axis led to the forma-
environment and to conduct the growing pollen tube tion of the conifer cone. By these transformations
to the megagametophyte for fertilization. As these the female reproductive organs were aggregated in
ovules were borne variously on branching systems, compound cones, which later again became reduced
leaves or other appendages, but otherwise exposed in various ways but the homology is often still dis-
to air and not enclosed in a capsule (ovary), like in cernible in ontogeny. The male reproductive organs
angiosperms, they were ‘naked’ as denoted by the (pollen cones) remained, or became early on, a sim-
term ‘gymnosperm’ which is thus merely a denomi- ple axis bearing microsporophylls with dehiscent
nator for a ‘less advanced’ mode of seed habit. All microsporangia (pollen sacs). Pollen cone morphol-
seed plants were by definition gymnosperms until ogy is rather uniform throughout the conifers. All
the angiosperm ‘seed habit’ evolved from a gymno- these (and other) transformations are well docu-
spermous seed. By sharing the evolutionary novelty mented in the fossil record. They can generally be
of having ovules and seeds enclosed in ­capsules, understood as adaptations to increasing aridity and
seasonality with cold winters, which indeed marked On its long journey from southern latitudes as a dis-
the end of the Carboniferous and beginning of the connected chunk of Gondwana, going through the
Permian in large sections of the supercontinent hot tropics and one or two arid zones it must have
Pangea. Out of perhaps a single ancestor (but which lost them all. On the collision with Asia, thrusting
one?) plants we now call conifers evolved, spread up the Himalayas, northern conifers spread south-
across much of the ancient supercontinent, were ward to occupy the mountains. India has but a single
dominant almost everywhere when this was divided southern conifer today: Nageia wallichiana, which
by the Thetys Ocean into Gondwana and Laurasia, is in fact an Indo-Malesian element spread from
and prevailed even when these two continents broke Southeast Asia. Another remarkable case is Ant-
up and the present continents started to drift further arctica. This continent has been glaciated for more
and further apart. But then, in the early Cretaceous, than 20 million years, perhaps completely for 15 mil- 15
the first angiosperms evolved and spread to all con- lion years. But prior to that it was vegetated and in
tinents eventually dominating most of the ecosys- the Mesozoic is was a major centre for plant evolu-
tems suitable for vascular plants. tion, the pivotal piece in the Gondwanan puzzle of
a fragmented continent of which the pieces started
In the Late Carboniferous and Permian a group of to drift apart just when ‘modern’ conifers evolved.
conifers known as Voltziales or ‘Walchian’ conifers Palaeobotanists are discovering more and more fos-
was widespread in Laurasia. Several had foliage sil sites in Antarctica and other Gondwanan conti-
leaves remarkably similar to some extant conifers, nents, yielding important remains especially for the
but the ovuliferous cones and especially the seed- understanding of southern hemisphere taxa. A third
bearing structures in these cones were very dif- phenomenon is the discovery of diverse conifer for-
ferent. The famous studies of Rudolf Florin in the ests that existed at very high Arctic latitudes such
1940’s and 1950’s solved once and for all the century as Axel Heiberg Island in Canada and Spitsbergen
old debate over the homology of the conifer cone from the Cretaceous into the early Tertiary where
and demonstrated how the conifer cone has evolved, now even tundra plants are few and very small. In a
at least in broad terms, from these Permian ances- warm temperate climate there existed mixed conifer
tors to, e.g., present day pines. ‘Modern’ conifers forests reminiscent of those occurring in, e.g., China
appear in the Triassic with Podocarpaceae and Arau- today, but which had to adapt to four or five months
cariaceae, of which some remains show traits that without daylight in winter. Many of these conifers
did hardly or not at all change during all the ensuing were the deciduous ones we know today from far
millions of years, for instance the structure of coni- more southern latitudes, such as Metasequoia, Taxo-
fer wood. Conifer wood is well preserved in numer- dium, Pseudolarix, and even Larix; the latter does
ous permineralized fossils allowing detailed study of occur in part within the Arctic circle today.
cell walls. The wood of many conifers is anatomic­ally
relatively simple in structure and often hardly distin- Conifer phylogenetic systematics is marked by one
guishable between families. The famous remains of predominant factor: extinction. Even the extant
Late Triassic (ca. 210 million years ago) conifer tree conifers reflect this very clearly, with 30 genera
trunks of the Petrified Forest in Arizona, USA could (43%) represented by just a single species and a fur-
have belonged to Araucariaceae, Podo­carpaceae, ther 11 genera with only two or three species, com-
Cupressaceae (Taxodiaceae) or more likely an ex- prising 59% of the total of genera. Further analysis
tinct family. The big logs of these Triassic trees were of their relationships, as well as the fossil record, are
transported by ancient rivers from a considerable strong indicators that this lack of diversity at spe-
distance upland where they once grew to the place cies level is the result of extinction, not a reflection
where they now lay. Yet, these and other fossils dem- of incipient speciation. Take for example the Dawn
onstrate abundantly how important and widespread Redwood (Metasequoia glyptostroboides), first found
conifers have been throughout the Meso­zoic. Of as a Miocene fossil in Japan, then as a very restricted
particular interest is the conifer history on the In- living tree in China, and subsequently (described
dian subcontinent, which is now almost devoid of as several species) from Late Cretaceous and Terti-
conifers (with exception of the Himalayan region ary deposits across the northern hemisphere even
which had a different geological history), but rich as far north as the arctic islands of Canada. In
in conifer fossils and those of other gymnosperms. 1994, a conifer recognized as a third living genus
 in ­Araucariaceae, Wollemia nobilis, was discovered molecular data derived from organisms that only
some 100 km northwest of Sydney in a remote, deep represent hugely disparate lineages due to extinc-
canyon; it has almost certainly near relatives in the tion. These difficulties will be only partly overcome
fossil record of Australia and beyond, but no more with more comprehensive sampling of extant taxa
than 100 mature trees survive ‘miraculously’ in the (Mathews, 2009). The inclusion of Gnetales in Coni­
wild today. Similar stories abound in conifers, and ferales (Pinales in Mabberley, 2008; Pinophyta of
while the popular press hailed both examples as ‘liv- other authors) is, therefore, premature at best and
ing fossils’ when they were first discovered, that label is not accepted in this book. Nearly all recent mo-
would be appropriate to scores of other species. The lecular analyses of the phylogenetic relationships
fossil record of conifers is comparatively rich and among extant conifers divide them into two clades:
16 very diverse, with numerous extinct species (Stew- the Pinaceae in one clade and the remainder of the
art & Rothwell, 1993; Anderson et al., 2007). Once families in the other (‘Cupressophyta’). Araucaria­
important groups or families have become extinct ceae and Podocarpaceae are placed in the latter clade
long ago, others are at present merely represented as sister groups (e.g. Rai et al., 2008). In the fossil
by a few scattered species, but some, even though record, however, these two and not Pinaceae are old-
they evolved long ago in the Mesozoic, are still with est (Farjon, 2008) and so we have again a conflict of
us and thriving. Although Florin (1951) considered evidence, albeit less serious than the ‘gnepine’ con-
the Carboniferous Cordaitales to be ancestral to troversy. Without consideration of the fossil record
conifers based on demonstrated homologies, more the relationships of the few remaining families of
recent analyses have cast doubts on this hypothesis conifers are of only limited ‘academic’ interest.
(Rothwell & Serbet, 1994; Rothwell & Mapes, 2001).
Modern cladistic analyses based on molecular data The 615 species of extant conifers are classified in 8
(DNA) assessing extant gymnosperms usually con- families of which 541 belong to the three largest fam-
clude on the monophyly of conifers (e.g. Stefanović ilies Pinaceae (232), Podocarpaceae (174) and Cupres-
et al., 1998; Rai et al., 2008), but these studies often saceae (135). Conifers are here considered to include
suffer from undersampling of taxa. As a conse- Cephalotaxaceae and Taxaceae; the Taxodiaceae are
quence of this, some extant taxa have been found to subsumed in the Cupressaceae with the exception of
be closely related using molecular data (not available Sciadopitys (‘Japanese umbrella pine’) which is as-
in fossils) while they appeared unrelated when mor- signed to its own monospecific family. Then there is
phological data from the fossil record were included the ancient family Araucariaceae, since the discov-
with those from living taxa. The most controversial ery of Wollemia with three genera, and finally the
case of this lack of agreement in cladistic analyses of still somewhat debatable family Phyllocladaceae (by
diverse data sets involves the conifers and a group some included in Podocarpaceae). Within the spe-
of gymnosperms known as Gnetales. The molecu- cies, subspecies and varieties are also recognized,
lar evidence often indicates a close relationship of which brings the total to around 800 different coni-
conifers and Ephedra, Gnetum and Welwitschia, the fers living today. Although such a figure is not fixed,
three rather disparate and relict genera constituting it gives a plausible estimate of the diversity at this
the extant order Gnetales. Despite seriously conflict- level. As has already been hinted at above, behind
ing results among several of these studies (e.g. Rai this rather moderate number of taxa lies a biological
et al., 2008; see for this conclusion also Mathews, diversity far greater than in any other natural group
2009), some analysts have boldly stated: “Gneto­ of plants of comparable size. A long evolutionary
phytes are derived conifers” and ‘found’ them to history and ecological marginalization by the gener-
be a sister group to Pinaceae (e.g. Hajibabaei et al., ally more competitive angiosperms since the mid-
2006). Such conclusions are superficial because they dle Cretaceous have led to numerous adaptations to
ignore the fossil record and thereby crucial evolu- cope with extreme environments, some examples
tionary information (Farjon, 2007). Indeed, cladis- will be mentioned under the brief outlines of fami-
tic analyses based on morphology that sampled the lies below.
fossil record did not give these results (Crane, 1985
and all subsequent studies) and appear to be much The Pinaceae, with 11 genera and 232 species are an
more consistent (Rothwell et al., 2009). There are exclusively northern hemisphere family. No fossils
perhaps analytical problems inherent in the use of are known from the southern hemisphere or from
parts that once were situated south of the equator. cones open their hard scales not enough to release
They are the ubiquitous pines, spruces, firs and larch- the seeds either. Which are the dispersing birds? We
es of northern boreal forests and mountain forests don’t really know, but when I was once on the one
extending further south into cool to warm temper- mountain in the world where this pine occurs, I saw
ate climatic zones. The pines (113 species) exhibit the some sinister ravens hopping about… could they
greatest diversity within a genus of this family, both have been the ones helping the tree regenerate?
taxonomically and biologically. They range from Firs (Abies) are by comparison with pines much
‘creeping’ shrubby species like Pinus pumila from more alike, but this book recognizes 47 species.
NE Asian mountains with a cold maritime climate They are in many respects more ‘demanding’ eco-
and P. culminicola on a few isolated mountain sum- logically, needing better soils and growing under cli-
mits of NE Mexico to 60 or 80 meters tall, straight matic conditions with less moisture stress and more 17
pines like P. ayacahuite in Mexico and P. lambertiana equitable, cool temperatures. Hence, they have to be
in California and Oregon. They occur on the Arctic able to out-compete angiosperms, or be codominant
tree line of Canada and Russia (P. banksiana, P. syl- in mixed forest. The seed cone bearing branches are
vestris), with a winter of 8 months and –50 °C tem- situated in the top of the tree, where the beautiful
peratures, but also in steaming hot ‘pine savannas’ erect cones can freely receive windblown pollen and
on lowland coasts of the Caribbean and in Thailand later disperse scales and winged seeds to the winds
and Cambodia (P. caribaea, P. kesiya), where tropi- that sweep through the crowns. Firs are mountain
cal rainstorms in the wet season alternate with fre- trees, which are rarely extending to lowlands like in
quent grass fires in the dry season. Growth rates can the northern taiga and there they are restricted to
vary in pines as much as size. There are amazingly more favorable sites for tree growth than other local
fast growers, such as P. palustris in the warm SE of conifers. Spruces (Picea), with 38 species, are better
the United States, adding 2–3 m of height per year, adapted to acidic soils and harsh climate than firs,
and pines that grow so slow that annual increment although there are exceptions, such as ­Picea sitchen-
rings cannot be discerned without a magnifying sis which occurs within a narrow coastal strip with
lens. Some of the latter, notably P. longaeva, carry on much rainfall and moderate temperatures from
for millennia. The oldest still living specimen tree in Alaska to California. It reaches enormous sizes in
the world growing in the White Mountains of Cali- the area around Vancouver and Seattle. Spruces can
fornia is more than 4806 years old (Lanner, 2007). form dense, uniform forests over very large areas,
It is the tree’s minimum age, an unknown number a quality used to advantage in plantation forestry.
of years must be added for the young tree to have Larches (Larix) with 11 species are deciduous, light
attained about 1.3 m height. It was a young vigor- demanding pioneers that colonize areas after dis-
ous sapling of a few centuries when the pyramids turbance, or occupy the most extreme habitats in
of Gizeh were being built during the 4th Dynasty of peat bogs of the boreal zone or the Arctic tree line,
ancient Egypt. While most pines disperse their seeds where on the Taymir Peninsula in Siberia at 75° N
by dropping them from opening cones, to be carried Larix gmelinii is the northernmost tree in the world.
away by wind aided by wings, some have evolved a Other genera have a limited number of species, e.g.
strategy dependent on mutualism with birds which Pseudo­tsuga and Tsuga, or only one, like Pseudo­
carry the seeds to caches for winter storage, where larix, a true relict now very rare in China (and not
many of them germinate. There are even one pine- closely related to Larix) and the enigmatic Cathaya,
one bird relationships, such as P. cembra with the also in China.
Eurasian Nutcracker (Nucifraga caryocatactes) in
Europe and P. albicaulis with Clark’s Nutcracker The Podocarpaceae, by contrast, are a largely tropi-
(Nucifraga columbiana) in North America. Seeds of cal family, outside the tropics they occur mostly in
these pines are ‘wingless’ and will not fall out of the mountains of the southern hemisphere. At present,
cones but have to be pried out by a bird’s bill. Pine 18 genera are recognized, with the largest, Podo­
cones range enormously in size, from 2 cm to 60 cm carpus, with 97 species nearly as speciose as the
in length and weights of a few grams to more than pines (Pinus). Next is Dacrydium, with 22 species.
2 kg in the amazing species P. maximartinezii of Za- The other genera all have fewer than 10 species.
catecas, Mexico. This pine has a similar seed mor- The family is undoubtedly of Gondwanan origin,
phology as the known ‘bird pines’ and the enormous with fossils going back to the Triassic found on all
 southern land masses including Antarctica. In the a little raspberry. Few studies are known that have
Podocarpaceae, taxonomic research is far from com- dealt with the question of what animals act as dis-
plete. On the one hand, more detailed observations persers; looking at these studies and what is known
of morphology, now often backed-up by molecular from pines, it is obvious that in Podocarpaceae mu-
evidence, has led to the recognition of more genera tualisms are likely to abound and much is still to be
to accommodate species formerly classified un- discovered here. That is not easy, because so many
der Podo­carpus and to a lesser extend Dacrydium. podocarps are scattered in tropical montane forests
On the other, quite a number of species in Podo­ and there are so many species of bird and other fruit
carpus are only known from a few herbarium col- eaters around to be suspected! As in Pinaceae, di-
lections, often with insufficient material and little or versity of life forms is great in this family, not only
18 no knowledge of the biology of the plant. A com- ranging from dwarfs like Microcachrys to giants like
prehensive revision of this genus, and of the entire Podocarpus totara of New Zealand, but even includ-
family, is wanting. A critical taxonomic revision will ing a true parasite, the only one in gymnosperms.
most likely lead to a reduction of the number of spe- Parasitaxus usta, from New Caledonia, lives on
cies recognized, but there may also be new discover- the root bases of another podocarp, Falcatifolium
ies in under-collected regions, such as western New taxoides. Some species are shrubs well adapted to
Guinea and southern Venezuela. For this book, only nutrient-poor and harsh environments, such as the
a preliminary revision of the family was possible, so 3 species of Lepidothamnus, one of which grows in
some relatively recently described species have here peat bogs in southernmost South America, the two
been retained ‘with the benefit of the doubt’. The others in the mountains of New Zealand. There is
Podo­carpaceae are characterized by specialized seed also a true rheophyte, the dwarf tree Retrophyllum
cones, in which in most cases during ontogenesis the minus from New Caledonia, with thick spongy bark,
number of seeds is reduced to one, which becomes perhaps to protect it from rocks and other debris
variously enclosed or subtended with soft, often smashing against it in flash floods. Others are emer-
colourful and at any rate to certain animals tasteful gent trees in tropical montane rain forests, such as
tissue. As a result, these conifers are adapted to dis- many species of Podocarpus, or shrubs to dominant
persal of seeds by animals (zoochory). However, as trees in the moss-forests of high tropical mountains,
in other conifers, the Podocarpaceae are wind polli- like species of Dacrydium in New Guinea.
nated. Hence, the pollen cones of the Podocarpaceae
do not differ from types found in the Pinaceae, but The Cupressaceae are the only cosmopolitan family
their seed cones are profoundly different. With few of conifers at present. The family is currently recog-
exceptions, a compound cone structure, with an axis nized to have 30 genera and 135 species. This count
beset with bracts and axillary ovuliferous scales, can includes the former, well-known family Taxodia­ceae,
only be discerned in the initial (pre-fertilization) a family concept which cannot be maintained when
phase. As soon as the first ovule, e.g. in Podocarpus, all the information now available, including phylog-
has been fertilized, the other ovules are aborted and eny, is considered. The species accomodated in the
the bracts swell and fuse together to form a juicy and Taxodiaceae were in fact a loose assembly of relict
colourful (often red or purple) receptacle to attract taxa having retained a few ‘primitive’ traits, such
birds. The initial cup-like seed scale grows around as spiral arrangement of leaves. Metasequoia, with
the seed to form the epimatium, which in some decussate leaf arrangement (phyllotaxis) is by many
genera (Afrocarpus, Nageia, Prumnopitys) swells to other characters related to Glyptostrobus, ­Sequoia
a succulent ‘fruit’ with the same attraction, taking and Taxodium with spiral phyllotaxis. Decussate
over the function of the receptacle. When a bird and whorled phyllotaxis, as well as other leaf char-
eats the ‘fruit’, the seed is swallowed with the rest, acters, appear to have evolved more than once from
but is protected from digestion by a hard seed coat a presumably ancestral spiral arrangement. While in
and dropped somewhere away from the parent tree. the Podocarpaceae extreme reduction of parts has
In some species, like the diminutive Microcachrys led to a single-seeded ‘cone’, in Cupressa­ceae reduc-
tetragona, a creeping dwarf shrub from the high- tion of another kind has taken place. Here, the seed
lands of western Tasmania, most ovules develop af- scale itself has been lost or, in some cases, e.g. in
ter fertilization; the bracts of the cone swell up and Cryptomeria japonica, vestiges in the form of seem-
turn bright red so that a ripe seed cone resembles ingly functionless appendages of what may have
been a scale, remain. The erect ovules (most other evolved there more recently. Yet others, such as
conifers have inverted ovules) originate on the apex A. araucana and A. angustifolia from South Ameri-
of a shoot in the axils of transformed leaves (bracts). ca and A. bidwillii from Queensland, Australia, may
These bracts enlarge in various ways and ultimately have changed little since the Jurassic. In contrast
form the cone scales, covering the seeds. with Araucaria, Agathis, with 17 species, is restricted
The worldwide distribution of Cupressaceae is to Malesia, NE Australia and islands of the SW Pa-
reflected in the great ecological diversity among cific. It is not yet known in the fossil record prior to
its species, although only a few are tropical such as the Tertiary, nor from other land masses. The most
some species of Callitris and Libocedrus from north- famous member of this genus is the Kauri, Agathis
ern Australia and New Caledonia and Papuacedrus australis, of the northern part of North Island, New
papuana from New Guinea and the Moluccas. The Zealand. The third genus, Wollemia, was discovered 19
other species occupy all the major environments in 1994 in a hidden canyon in Wollemi National
that conifers thrive in generally, but in addition have Park, New South Wales, Australia, and is one of the
many species well adapted to semi-arid conditions. most interesting of recent conifer discoveries. It is
Especially diverse in semi-arid environments, which considered to be another ‘living fossil’ and true fos-
occupy large parts of the land masses of four conti- sils are indeed now being assigned to this new genus.
nents, are Cupressus and Juniperus in the northern
hemisphere and Callitris in Australia; some of the The Phyllocladaceae are distinguished by greatly
species even occur in true deserts. Other extremes reduced true leaves and ‘phylloclades’, representing
are altitude records, e.g. Juniperus indica found at planated shoots, taking on the tasks of assimilating
5100 m altitude on the moraines of Tibetan glaciers leaves. They also have a distinct system of pollen
coming from Mt. Everest. Other Cupressaceae thrive capture by the ovule, which sets them apart from
in ecosystems that can hardly be more different most, but not all podocarps, as does the formation of
than these: the wet, oceanic forests of giant conifers a true aril around the seed, as in the Taxaceae. How-
along the Pacific coast of North America from cen- ever, molecular evidence appears to indicate that
tral California to Alaska. Sequoia sempervirens from they are very closely related to the Podo­carpaceae.
California is the tallest tree in the world, reaching The Phyllocladaceae are from a Gondwanan ances-
up to 117 m. Sequoiadendron giganteum, confined to try and greatly reduced. The four species of its single
the western slopes of the Californian Sierra Nevada genus are now scattered in Malesia, New Zealand
a little further inland, is a living ‘fossil’ of gargantuan and Tasmania. The Sciadopityaceae are represent-
proportions. Compare this with the dwarfed species ed by a single species, Sciadopitys verticillata from
Microbiota decussata from the Russian Far East and a few localities in Japan. This conifer has until re-
Diselma archeri from Tasmania, both important cently been misunderstood; it is not a pine and al-
components of subalpine scrub communities, and though related to the Cupressaceae it has a whole
the diversity in this family, and that of the conifers suite of unique characters. It is a prime example of
in general, will become clear. a relict taxon and deserves more intensive study by
all disciplines relevant to its taxonomy. The Cephalo-
The smaller families, except the Taxaceae, have a taxaceae, with a single genus as recognized by most
more limited distribution. The Araucariaceae occur taxonomists, accommodates 8 species, all eastern
in South America, with two species, one of which Asian, ranging from Korea and Japan to Malaysia.
is the well known ‘Monkey puzzle’ Araucaria arau- In some DNA-based analyses, this family has been
cana, but is more abundant in Borneo, New Guin- proposed for synonymy with Taxaceae, but I think
ea, NE Australia, and New Caledonia and other there are arguments to retain it as a separate fam-
SW Pacific islands. The genus Araucaria was in the ily. Several species are cultivated widely and the tax-
Mesozoic nearly cosmopolitan, and hence the typi- onomy is still somewhat unsettled, in part because
cal conifer with which to depict the dinosaurs, but some species are based on cultivated plants, in part
became increasingly restricted in its distribution. It because species have been based on variable foliage
does not mean that all the 19 species currently rec- characters due to the extremely reduced ovulifer-
ognized are ‘living fossils’; it is more likely that most ous cones. A similar difficulty besets the Taxaceae,
of the species in New Caledonia, where at present and especially its largest genus Taxus, here recog-
the genus has its centre of diversity with 13 species, nized with 10 species, but recently again the subject
 of ­debate, when a long-time student of the genus it and more fleshy and bluish green to purple. The
raised the number of species to 24, with 55 varieties aril is eaten by birds which disperse the seeds un-
(Spjut, 2007). In the Taxaceae, 4 other genera are damaged. In recent years, the discovery of chemi-
recognized, of which Amentotaxus and Torreya have cal compounds (taxanes), which have proven to be
6 species. All are evergreen shrubs or small trees in certain cases effective against cancer in humans,
adapted to live in the understorey of tall forests and have focused a lot of research on the species of Tax-
have developed a single arillous seed of which the us. This event has also demonstrated how much we
aril is either partly covering the seed and soft and can still learn about conifers.
juicy and colorful red or yellow, or wholly covering

20
THE DISTRIBUTION AND ECOLOGY OF CONIFERS
The 615 species of conifers cover a large proportion while the Greenland icecap prevents any form of
of the land surface of the earth. The map shows that vegetation. The steppes of North America, Central
the greatest covering of land by conifers is clearly to Asia, Tibet and Mongolia, parts of Africa, and Pat-
be seen in the northern hemisphere. This is due to agonia in South America can have some conifers
the extensive conifer forest of the boreal zone in Eur- here and there, but are also largely without them. Al- 21
asia and North America. There are only few species though a few conifers occur in deserts, most of the
occurring there and the diversity increases dramati- world’s deserts, i.e. the deserts of central and western
cally further south, while the areas being covered Asia, Arabia, North Africa (Sahara), and Australia,
decrease. The area indicated to be covered by coni- are devoid of conifers. Another vegetation type
fers on the map is, of course, not uniformly covered mostly devoid of conifers is lowland tropical rain-
with conifers, they usually occur together with an- forest, such as the Amazon Basin in South America
giosperms or grow in patches too small to separate and the Congo Basin in Africa but also in smaller
on this scale. There are already big gaps at around areas like the Yucatán Peninsula in Mexico and the
30° North, which continue across the equator into southern lowlands of New Guinea. Other large areas
the southern hemisphere. In the far south there is without conifers have been stripped of their natural
much less land, but except Antarctica – which has vegetation and are now occupied by agriculture and
been omitted from the map for obvious reasons – it urbanization, especially in Western Europe, India,
is quite well covered with conifers, with few big gaps. Bangladesh and northern China. However, some
Conifers are present in nearly all the major veg- of the remaining areas devoid of conifers cannot be
etation types of the world. They are absent from only resolved with an ecological explanation. Large parts
a few and are very rare in some others. In the far of South America and larger parts of Africa and the
north, a few conifers occur in tundra vegetation, Indian subcontinent are the major gaps in coni-
mostly in the transition zone from boreal forest to fer distribution that have no explanation in terms
tundra. In the High Arctic of northern Canada and of unsuitable climate or soils. Did these parts ever
northern Siberia (cold deserts) they are left behind, have conifers, and if so, why did they loose them?

The global distribution of conifers shown on a world map with approximate equal area projection. The
black areas on the map are not uniformly covered with conifers; they occur together with angiosperms or in
patches too small to separate on this scale.
 ­ iscounting the Mediterranean coastal areas, which
D the Cupressaceae seem to throw the separation of
biogeographically belong to Eurasia, Africa is the extant conifers into northern and southern origins
poorest continent for conifers today. In Sub-Saharan into doubt. Araucaria may have originated before
Africa we only have Widdringtonia (Cupressaceae) the separation of Pangea into two super-continents,
with four species, Afrocarpus (Podocarpaceae) with but as far as we know, the Cupressaceae evolved after
five species and Podocarpus with four species. Vast that event.
areas of Sub-Saharan Africa have no conifers at all.
The Indian subcontinent south of the Himalaya and One final, intriguing observation about present-day
adjacent hills has only a single indigenous conifer: conifer distribution is that more than half (ca. 330)
Nageia wallichiana (Podocarpaceae). There are plen- of all species occur on the Pacific Rim. All families
22 ty of suitable areas for conifers in both, as the suc- and 83% of all 70 genera are represented. The Pacific
cessful plantation of conifers for forestry in Africa Rim is the system of mountain ranges and islands
and India demonstrates. The causes are most like- around the Pacific Ocean in both hemispheres,
ly of a historical kind, having had effect over time mostly forming part of so-called subduction zones
spans on a geological scale. It appears that isolation, where oceanic crust plates slide beneath the conti-
caused by the break-up of the southern superconti- nents, causing volcanism and thrusting up of moun-
nent Gondwana and the increasing separation of its tains and islands in the process. Going clock-wise an
constituent land masses, is a major factor behind the starting at 10 o’clock the diversity centres for coni-
gaps in conifer distribution of land masses of Gond- fers are Japan, the Pacific North-west of the United
wanan origin. Connections between land masses re- States and Vancouver Island, California, southern
mained much longer in place with the break-up of Mexico and Guatemala, southern Chile, New Zea-
Laurasia, the northern super-continent, which also land, Tasmania, New Caledonia, Fiji, the coast of
became less fragmented. Both dispersal and vicari- New South Wales and Queensland, New Guinea,
ance played a part in the history of conifer distri- Borneo, the Philippines, and Taiwan. All of these
bution leading to the present situation. However, areas have many species in several genera and fami-
due to the size, proximity, and orientation of land lies. In contrast the coastal areas of the Atlantic and
masses of Laurasian origin, all situated in the north- ­Indian Oceans do not have such diversity of conifers,
ern hemisphere, dispersal accounts for much of the with the exception of Morocco, which has a moder-
distribution of genera and species. Losses in the past ate diversity of 12 species, and Florida, with 10. The
could often still be made up for by new arrivals. explanation is probably again historical: none of the
The long history of the assemblage and subse- other oceans are nearly as old as the Pacific Ocean,
quent fragmentation of the two super-continents, which has therefore provided coastal mountains and
Laurasia in the north and Gondwana in the south, is islands suitable for well moisturized montane forests
apparently still reflected in the distribution patterns from well before the origin of the angiosperms.
of conifers across the globe. If we look at the differ-
ent taxa this is even more clearly demonstrated. At In the north, around the Arctic Circle, the conifer
the family level, Cephalotaxaceae, Pinaceae, Sciado­ forest or taiga begins where the tundra ends. The
pityaceae and Taxaceae are Laur­asian, whereas Arau- taiga is not entirely homogeneous. It is interrupted
cariaceae, Phyllocladaceae and Podocarpaceae are by lakes and swamps and dissected by large rivers.
Gondwanan, although Araucaria once extended into There are few conifer species and most of them be-
Laurasia. Cupressaceae as a family are cosmopolitan, long to the family Pinaceae. In some areas two or
but the 30 genera that make up this family today three of these conifer species may occur together, but
are divided in northern and southern hemisphere very often the forest is formed by only a single spe-
groups, with a few ‘trespassers’, some only known cies. The environment of these conifers is dynamic.
from the fossil record. Some of these ­exceptions Disturbance by fire and flooding is an integral part of
are intriguing enough, though. The South African the taiga ecosystem, and natural disturbance, mostly
genus Widdringtonia has been found in the Upper in the form of fires, often covers large areas. Storms,
Cretaceous of North America and ­Austrosequoia however, are rare. The air burst of a large meteoroid
wintonensis from the Cretaceous of Australia is very or comet fragment in June 1908 in Siberia, known as
similar to Sequoia or Sequoiadendron of Laurasian the Tunguska Event, destroyed large tracts of taiga
origin. Such widely distant ­occurrences of taxa in forest. Outbursts of defoliating insect plagues, often
associated with freak weather conditions, can lay key factors for the success of conifers on poor soils
waste to the forest as well. However, usually within a must be the extensive development of mycorrhizal
few decades the conifer forest is coming back from symbiosis with certain fungi, which is especially
abundant seed. It was their capacity to conquer new prevalent in conifers. This form of collaboration be-
land in great numbers that enabled conifers to return tween entirely different organisms enables conifers
and to occupy the vast northern regions during the to cope with very low levels of essential minerals
relatively short interglacial phases between the ice such as phosphorus and nitrogen in the soil, because
ages of the Pleistocene. But the present conditions the fungal hyphae greatly expand the surface and
are still harsh, the growing season is very short, and thereby uptake of water and nutrients by the plant
only a few species have been able to adapt to these. root system. It is now even being thought that plants
would not have succeeded to come onto dry land 23
Conifers predominantly occur on soils or in cli- permanently without the assistance of these fungi;
mates that are sub-optimal for plant growth. How- conifers were among the first to succeed in leaving
ever, as we know well from plantation forestry and the lakes and swamps behind them.
horticulture, there is no evidence that conifers pre-
fer nutrient-poor soils or harsh climates. Many spe- Although it is the most widely applied strategy, not
cies that are restricted to these poor conditions in all conifers are escapists. Especially in areas on the
their natural habitat grow exceedingly well if plant- Pacific Rim, there are many conifers that grow very
ed on fertile soils. Monterey or Radiata pine (Pinus tall, rising well above the general forest canopy. Most
radiata) has been planted on a large scale especially of them can live very long, some even thousands of
in the southern hemisphere and is one of the most years. These giant trees appear to be scattered in the
productive trees in the industry. Yet its natural habi- forest as veteran trees, or they occur in cohorts of
tat is nutrient-deficient podzolic sand in the salty more or less even-aged trees when still of a lesser
winds of the Pacific Ocean right on the Californian age. These forests grow on the whole on mineral-
coast. Many species of conifer are more or less re- rich, young soils, often of volcanic and sometimes
stricted to ultramafic soils derived from serpentine of glacial origin. Competition among abundant
or similar rocks, where phosphorus is almost ab- trees of many species is fierce. The ecological strat-
sent. It appears that conifers, in order to survive in egy here is closely tied in with localized forest dis-
nature, have often adapted to such poor sites. It is turbance. Fires, land slides and destructive storms
the strategy of evasion. What are these conifers try- in these places occur infrequently, sometimes with
ing to escape from that they have adapted to put up intervals of several centuries. The old, big trees are
with such poor conditions? The answer is competi- the survivors of such disturbances. The bigger the
tion from angiosperms, in particular fast growing, tree, the more resistant it becomes; the bark of huge
large-leaved herbs, shrubs and trees. While conifers, trees of Sequoiadendron giganteum from the western
once they are full-grown trees, can often outgrow slopes of the Sierra Nevada in California is up to 60
angiosperm trees, they often have a slower start. It cm thick and a very effective protection against for-
has long been assumed that the evolution leading to est fires. Nearly all of these giants among the coni-
weedy herbaceous angiosperms has caused the de- fers are light demanding and would not successfully
cline of conifers. Grasses are particularly effective regenerate under the canopy of other trees, being
in suppressing juvenile conifers and have excluded either angiosperms or conifers. These trees have to
conifers from the steppes in Asia and the Ameri- be removed first. On open sites, regeneration of the
cas and probably from the grasslands of Africa. big tree conifers is often abundant, but soon compe-
That conifers can grow well there is again demon- tition starts and a struggle for space, light and ­water
strated by plantation forestry. Overgrazing and fire begins. Since some of the big conifers can grow tall-
prevention, suppressing the grasses, can lead to the er, they have a chance to become dominant above
return of conifers; this happens at present with juni- the canopy of the other trees. The giant ultimate sur-
pers (Juniperus) on the rangelands of the American vivors are the insurance policy of the species.
West. However, if site conditions are poor for any
plant growth, the conifers have a natural chance. There are many mutually beneficial relationships be-
Once they are mature, they will persist because they tween animals and plants, almost all of which have
now can compete with any angiosperm. One of the developed with angiosperms on the plant’s side of
 the deal. Pollination provides numerous examples saceae, it is the genus Juniperus in which the cone, in
and it is the single reason why flowering plants de- a very different way, has become fruit-like. Birds are
veloped their almost endless variation of flower de- again the main dispersers of the seeds, in many spe-
signs. Not a single conifer has ever managed this as cies reduced to one large seed per cone; quite a re-
far as we know, they are all wind pollinated, which is, duction in number if the evidence is correct that the
lacking precision in pollen delivery, a wasteful pro­ genus may have arisen from an ancestral species of
cess. Wind pollination has, however, not ­prevented Cupressus, of which many species have well over 100
them from becoming abundant and widespread, seeds per cone. The seeds of all species in the Tax-
and some wind pollinated angiosperms, like grass- aceae arise at the apex of dwarf shoots, a seed cone
es, have done well, too. Many conifer species have does not even develop to become ‘aborted’ in a later
24 been more successful with seed dispersal aided by stage as in some species of the Podocarpaceae. An
animals. In general, there are two scenarios for seed aril, arising from the initial seed integument, takes
dispersal: (i) the animal transports and eats a fruit on the fruit function. The imitation of angiosperm
containing seeds, which pass through the animal fruits has proved very successful in dispersing the
undigested, and (ii) the seeds are transported and seeds of conifers with a minimal waste of resources.
hidden to be eaten later and not all hidden seeds are In the second scenario of seed dispersal by animals,
eaten. Conifers have adapted to both scenarios and some species of pine (Pinus) have developed mu-
the first one is more common than the second. The tualistic relationships with birds of the crow family
families Podocarpaceae, Taxaceae, Phyllocladaceae, (Corvidae) to effect seed dispersal. The birds remove
Cephalotaxaceae and Cupressaceae all have many the seeds from the specially adapted cones and hide
species in which the seed cone, or the seed, has them in the surrounding terrain for later consump-
evolved to resemble a fruit. In all cases, the cone has tion. Large quantities of seeds are involved and in-
been extremely reduced, in some cases it effectively variably not all seeds get eaten. Where their close
disappeared. With few exceptions, the number of relatives have winged seeds, the seeds of these pines
seeds has been reduced to one only. In many ways, are wingless; wings would be useless and a waste to
the Podocarpaceae, the second largest family of produce. In this way, the pines can travel upslope in-
conifers, has been the most innovative as well as the dependent of prevailing wind conditions, which for
most successful family to adopt this strategy of seed some of the subalpine species may turn out to be an
dispersal. Probably all 174 extant species in the fami- advantage during a warming climate.
ly have their seeds dispersed by birds. In the Cupres-
THE ECONOMIC IMPORTANCE OF CONIFERS
The economic uses of conifers can be conveniently ty with properties such as high dimensional stability,
divided in those centring on wood, other derivative straight grain, softness and workability together with
products, and horticulture. The first two are mostly large, straight dimensions especially when ­taken
industrial. Horticulture does not generally manu- from ‘old growth’ forests. These timbers are applied
facture products but focuses on the cultivation and in the building industry for, e.g. doors and win- 25
trade of the whole plant and has a strong element of dows, in furniture making and even for musical in-
non-commercial interest. However, there are mani- struments such as organ pipes and piano keys. Firs
festations of both applications evident in all three (Abies) produce lightweight, relatively soft, creamy
categories of economic use of conifers, so that the white to pale brown wood; the high grade timber
demarcation between industry and leisure is not an from forests with these species is sawn for framing
absolute one. It could even be argued that these dis- material and for plywood and veneer. The wood of
tinctions have become less clear in recent times than spruces (Picea) differs markedly from that of firs
they were in the past, and examples will be given in and is consequently used for different, mostly less
this chapter. refined purposes. In northern lands, it was and of-
Today the world trade in coniferous wood is ten still is the principal tree for the construction of
huge. Conifers provide about 60% of all wood used log houses. In Norway, the ancient village church-
for industrial purposes. They dominate industrial es were entirely made of wood, often a mixture of
wood supply because of both technical and econom- spruce (Picea abies) and pine (Pinus sylvestris); some
ic advantages over wood from angiosperms. The date from nearly 1000 years ago and are still intact.
vast reserves of natural conifer forest in the boreal In the Alps similar uses in construction are made
zone of the northern hemisphere play an important of larch (Larix decidua), as its wood is particularly
part in this supply, but plantations, especially in the resistant to weather and rot. All three genera also
southern hemisphere, of particularly pines take up provide pulpwood for the paper industry; the trans-
an increasing proportion of this. Nearly all this mass parent windows in envelopes are a paper made from
production goes into pulpwood for the paper indus- larch wood. In some genera of conifers, various cir-
try, although about ⅔ of conifers in plantations in the cumstances have caused only one of the species to
world are destined for timber. The more specialized, become of high economic importance. An example
high quality woods used for all the versatile applica- is the Douglas fir (Pseudotsuga menziesii), which has
tions mentioned in this chapter come from forests become the most important timber tree in North
that can be allowed to grow older. In the trade, the America. It has consequently been introduced by
wood of conifers is known as softwood and that of foresters in many temperate regions around the
angiosperms as hardwood. These terms are mislead- world. Douglas fir grows rapidly, is straight and tall
ing if applied to all conifers, because many species and produces large volumes of wood per hectare,
produce very hard wood indeed, while some of the especially in managed forests where competing spe-
softest woods in use do not come from conifers but cies are excluded or suppressed. Its wood is used
from angiosperms. By far the greatest volume of for plywood and construction, both exterior and
industrial wood is produced by species in the fam- interior, and it has a reasonable durability for out-
ily Pinaceae, which occur naturally in the northern door applications such as telephone poles and rail-
hemisphere only, but have been widely planted for way sleepers. The wood of the Cupressaceae differs
this purpose in the southern hemisphere. In par- markedly from that of Pinaceae. It is more fibrous
ticular, the genus Pinus stands out, with several and contains less resin; it is also mostly very decay-
species being among the fastest growing plantation resistant and many species have fragrant properties
trees producing pulpwood on short-term rotations. due to volatile chemical compounds. These prop-
Several other species, especially ‘white pines’ (Pinus erties make it highly desirable in China and Japan,
subsect. Strobi), are providing wood of higher quali- where it was traditionally used in the construction
 of temples and other ceremonial buildings. Some (Southern Africa), Dacrydium and Dacry­carpus
species have been over-exploited and good sized (Australasia) and Nageia (Southeast Asia). All
trees are now rare. Other species, like Cryptomeria yield light to medium-weight, pale coloured wood,
japonica, have been widely planted, in plantations in known as podo in the trade, with a straight grain
Japan, where it is native, as well as in China and Tai- and even texture that is easy to saw and plane but is
wan. Some of this cupressaceceous wood splits eas- often brittle. It is not durable when exposed to the
ily into shingles, and its rot-resistance was noted by weather, so its building applications are for indoor
people in regions as far apart as the Pacific coasts of construction only. High grade timber can be used
Canada and the northwestern United States, Chile, for door and window frames, panelling, veneer, cup-
Japan and Vietnam, where these shingles were tradi- boards, furniture, cabinet work, joinery, household
26 tionally used to cover the roofs of houses. Durability utensils and engineering instruments like drawing
has also been the major property that made juniper boards and rulers. The New Zealand species Podo-
(Juniperus) the tree of preference for fence posts, but carpus totara provides the only softwood that is re-
metal is pushing it out of the market. Some species sistant to attack by marine borers, so it was used for
in the Cupressaceae produce wood with beautiful ship and boat building as well as wharf building and
patterns and are therefore prized for the making of harbour construction until a ban on further logging
cabinets and other pieces of furniture. In particular, stopped it. The Maori built their famously long war
the large, ancient coppice stools of the North African canoes with the wood of this large indigenous tree.
species Tetraclinis articulata are valuable and were
already sought after by the Romans. Few conifers Of second industrial importance to wood of coni-
have this coppicing capacity, i.e. re-growth from a fers is their resin. Resin is present in all conifers, al-
stem base after repeated cutting. Another source for beit not in equal quantities. Resin in leaves can be
this type of wood, suitable also for wood turning, is distilled from them, resin in wood can be tapped as
yew (Taxus), which is hard, dense, heavy and resist- well as distilled, and there is even resin to be mined.
ant to decay. Perhaps most famous were the English The resins of conifers are mostly terpenoid, with
yew longbows of the Middle Ages; the arrows shot some phenolic resins (Langenheim, 2003). Only two
from these bows could penetrate a knight’s armour families, Araucariaceae and Pinaceae, produce copi-
at 200 meters and helped the English win the bat- ous amounts in the wood, where the resin is stored
tles of Crécy, Poitiers and Agincourt in the Hundred in resin ducts or canals. The genus Agathis (Arau-
Years’ War (1337–1453). cariaceae) is the most copious producer of resin;
The most tropical of the conifer genera, Agathis over centuries, resin has flowed from the trunks of
(Araucariaceae) is one of the most valued timber large trees onto the ground where it has accumu-
trees in Australasia. The wood known as kauri in the lated, forming large deposits of copal which can
timber trade is light and soft, pale yellow or straw- be excavated. Its use is mostly for varnishes. More
coloured, often with darker heartwood ranging commonly, resin is tapped from the trunks of pines.
from pink to dark red brown. The wood of Agathis Major resin producing species are, or were, Pinus
has many uses, such as indoor construction, panel- kesiya in Southeast Asia, P. massoniana in China,
ling, boat masts, joinery, furniture, pencils, matches P. pinaster in Europe, especially France, and P. palus-
and matchboxes, rulers, and piano parts. Naturally tris and P. elliottii in the Southeastern United States.
it is excellent for plywood and veneer, while more The resin tapped forms the basis of many products
industrial uses of lower grade wood are pulp for pa- in industry, such as turpentine, rosin and pitch – to-
per manufacturing and high grade charcoal. In In- gether known as naval stores – oils, varnishes, print-
donesia and Malaysia Agathis is exploited heavily for ing inks, sealing wax, soap, plastics, and fireworks.
export of raw timber (round logs); in the Philippines Coarser products are obtained by destructive distil-
this has already led to a total ban on further cutting, lation of resinous wood. In the age of wooden ships,
while export is banned from Papua New Guinea. In pitch and tar were indispensable to keep them sea-
tropical countries the wood of the Podocarpaceae worthy by caulking the seams with pitch and by tar-
is usually highly valued and trees belonging to this ring the rigging. The term naval stores for these and
family are often selectively logged for timber. The similar products dates back to the 17th century, when
most important genus, Podocarpus, is also the most the English navy required large quantities of these
widespread; of regional importance are Afro­carpus for an expanding fleet.
The value of conifers for human food is marginal Conifers in horticulture certainly started as a pas-
and consists almost exclusively of the seeds of sev- time, not as a business. Its earliest roots lie undoubt-
eral species of pine (Pinus) and three species of edly in China and Japan, where planting of trees for
Arau­caria, two in South America and one in Aus- aesthetic reasons antedates that activity in ­Europe
tralia. Pine seeds are nutritious and sometimes taste- by centuries. In that tradition, form, shape and
ful food and some are of commercial value today. colour were of greatest importance as part of a re-
In ­Europe, it is mainly the Mediterranean species construction in situ of idealized pictorial land- and
P. pinea or umbrella pine which yields seeds of good townscapes. By contrast, in Europe the planting of
size and taste for the food market (Italian ‘pigno- (exotic) conifers in gardens and parks mostly re-
lia’), used in processed food as well as sold as whole sulted from a curiosity about the natural world of
seeds. The distantly related Asian pine P. ­koraiensis distant lands and the products these brought forth. 27
produces similar kernels and has become the leading This developed in the 19th century to a rage among
species in the export market. Pinus gerardiana, or the landed gentry of Europe to obtain species, which
chilgoza, and P. bungeana, or lace-bark pine, are oth- led them to employ ‘plant hunters’ to travel to Amer-
er Asian pines with good edible seeds. In the United ica and Asia to obtain suitable tree seeds that could
States and Mexico there are several species with be grown in Europe. Horticulture developed in part
edible seeds, of which P. cembroides, P. edulis and from the demand that this trade generated, with
P. monophylla are the most widespread, which are nurseries in e.g. England and Germany specializing
commonly known as piñon or pinyon pines. In Rus- in trees, including conifers, for large gardens, parks
sia, P. sibirica produces seeds which are harvested for and estates. This again fed into plantation forestry,
oil extraction. The seeds of Araucaria, although deli- when some land owners saw potentials for planting
cious when roasted, are more of traditional value to timber trees on their estates. By the middle of the
indigenous people than commercialized for the food 19th century, conifers from North America were well
or delicatessen markets. The rarity of the trees of established and common, while the first introduc-
A. araucana and A. bidwillii due t o past over-exploi- tions from China and Japan had just arrived. About
tation prevents large scale seed harvesting. Flavour- 60–70 years later, these had become common, too. It
ing is probably best known from the use of juniper was this widespread planting of exotic conifers and
‘berries’ (seed cones) in producing gin. The Dutch other trees which caused a demand for descriptive
words for gin and for the shrub Juniperus communis literature and gave rise to a number of ‘manuals’ or
are, respectively, ‘jenever’ and ‘jeneverbes’, in the lat- ‘handbooks’, particularly in England and Germany.
ter the Dutch word for berry is added. This species, Naturally, these books concentrated on the species,
together with its varieties, is the most widespread and soon also cultivars, that were grown in Europe.
conifer in the world and the various brands of gin, The British Isles, due to a relatively mild but var-
like aquavit, gin, jenever and Schnaps, are all made ied climate and varied geology and topography, as
with it. The resin of the Mediterranean pine Pinus well as widespread private landownership that has
halepensis is used to flavour retsina, a Greek white continued to the present, are the world’s ‘centre of
or rosé wine popular with locals as well as tourists. diversity’ for introduced conifers. Current member-
Scent is close to flavour. The typical fragrance of a ship of the International Dendrology Society (which
northern conifer forest is due to the volatile com- was established on the continent) reflects this. How-
ponents of the terpenoid resins of the conifers and ever, in recent decades other parts of the world have
these are being used in the cosmetics industry. The been catching up, such as the United States, Aus-
volatile components were and are also used in re- tralia and New Zealand. With parts of these coun-
ligious ceremonies, as fragrance and scent play a tries in a warmer and either wetter or drier climate
major role in imagination. The Aztecs called teocote than most of Europe, the range of species that can
pine (Pinus teocote) the pine of the gods; to burn its be grown extends beyond the more or less common
rosin as incense was a privilege of priests and kings. assortment used in Europe. There is no good reason
Religious or symbolic use also lies at the root of the why tropical conifers should not be planted in gar-
Christmas tradition to decorate a room in the house dens and parks of tropical countries, which is one
with an evergreen tree and the growing of several reason why this book includes all species. While the
species of conifer for this purpose has developed in planting of conifers in gardens and private parks re-
the last 150 years or so to a significant industry. mains ‘pastime’ activity, horticulture with ­conifers
 has ­developed into a substantial business aiming achieve in this métier today. The trade is, of course,
at a different and wider market. In recent years the influenced by the demand but also creates it by mak-
trade appears to have been emphasising dwarf coni- ing new and better cultivars available. Many good
fers, but the fashion is not new. One of the many dwarf and colour cultivars have been developed in
garden crazes in the Victorian age was building rock the last decade or so. A major development has been
gardens in parks with plantings in it that ought to the utilization of witches’ brooms occurring as a
grow slow and remain moderate in size. In Britain, result of DNA mutations in buds or shoots. Unlike
dwarf conifers became popular for a time from the those resulting from a tree’s reaction to parasites or
1850’s onward. Interest was renewed in the 1960’s pathogens, these genetic mutations alter growth of
but this time it was a fashion to plant heathers in any shoot taken from the witches’ broom, resulting
28 suburban gardens and it affected Europe as well as in clones with identical, permanent dwarf growth
the United States. Consequently, many arboreta and characteristics. Although they were first used as
pineta followed suit and established heather gardens long ago as 1836, in recent times selections derived
with small or medium size, preferably columnar or from witches’ brooms have become a major source
fastigiate conifers. When people began to be bored of dwarf conifers in cultivation, and a constant
with this, the heather garden went out of fashion stream of new cultivars is entering the market. With
again, although it may still have its adherents. One a growing knowledge of causes and processes, espe-
reason for the disillusion may have been that ‘dwarf ’ cially genetics, and improved techniques in propa-
conifers often grow bigger after a slow start than gation and cultivation, we can see this develop in
they were promised to do when planted. The meth- future into an industry where conifer cultivars are
ods and practices of growing dwarfs only slowly being ‘made’ as well as grown.
developed into what professional nurserymen can
THE CONSERVATION OF CONIFER DIVERSITY
Extinction is an apparent fate of all species. What Against this rather overwhelming perspective, it
concerns conservationists is the current high rate of could appear to be next to hopeless to take action
extinction, well above what is believed to be a ‘back- for the conservation of conifer diversity in the natu-
ground’ rate. Different estimates of these rates exist ral habitats in which they occur. The real big issue
and are debated, but what is undisputed is that the is human demography, but to address that is both 29
current rate is abnormally high and that humans are complicated and difficult. It has to be addressed, or
causing it. “The primary cause of the decay of organ- else the size of the population will be ‘corrected’ nat-
ic diversity is not direct human exploitation or ma- urally, which will be disastrous. Meanwhile, we can
levolence, but the habitat destruction that inevitably and should buy time, although there is not much
results from the expansion of human populations time left. This is why the conservation of species in
and human activities.” This is how Paul Ehrlich, a their natural habitats is very important, and even
well known American ecologist and writer on the more urgent when certain species are ‘keystone’ in
biodiversity crisis, has summed up the cause of the the ecosystem. Many conifers perform that role,
present mass extinction of species (Ehrlich & Ehr- because they are large, long living trees providing
lich, 1981). He called the destruction inevitable and habitat for numerous other species. Conservation
that is what it is under those circumstances. He later of these conifer species in functioning ecosystems
calculated that by the end of the 1980’s humans had in the wild means the conservation of many other
consumed, directly and indirectly, up to 40% of the species. It is therefore a good strategy to concentrate
total net primary production of stored biological en- conservation efforts on conifers. Especially in the
ergy produced by plants from solar energy. So, one tropics, however, our knowledge of the role of coni-
species is using two fifths of all biological produc- fers in ecosystems is rudimentary. Where natural,
tion on the planet. He warned that when the hu- undisturbed forests are formed by many tree species,
man population doubles again, that could become conifers often occupy restricted localities and occur
as much as 80%. Logically, if human activities and there in numbers. The ecosystem will be different in
economic growth also continue to rise, we will one such places, in which the conifers are likely to act as
day have to use it all, and there will be no room for keystone species. Research is urgently needed where
vegetation not in the service of humanity. If every- logging and deforestation threaten the existence of
one in the world today enjoyed a lifestyle similar to such tropical conifers with local abundancy, e.g. the
the one people have in Europe and the United States species of Agathis in Borneo and Afrocarpus in east-
we would need at least three planets to sustain our- ern Africa. Conifers are also important in nature
selves; with a world population double that of today, because all trees are providers of ecological services
we would need six planets. Edward Wilson, a lead- to a more stable environment. Climate regulation is
ing writer on evolution and extinction, has put the one; watershed protection preventing erosion and
ecological absurdity of our numbers succinctly, not- assuring a steady supply of clean water in streams
ing that we are now 100 times more numerous than are other important functions of forests. In valleys
the most abundant species of large animals that has with human habitation, the prevention or limitation
ever existed. “There is no way that we can draw upon of the effects of snow avalanches in mountainous
the resources of the planet to such a degree without areas is obviously important. On slopes where ava-
drastically reducing the state of most other species.” lanche stopping conifer forests have been removed
(Wilson, 1992). That state for most species means it is nearly impossible to restore them and in popu-
extinction. The prospect that within a few decades lated valleys artificial avalanche barriers will have to
far more people will live in cities than in the coun- be erected. They are expensive and not as effective
tryside (the 50–50 balance line has recently been as the natural conifer forests. That conifers provide
crossed) threatens to detach humanity further from these services in many parts of the world is the logi-
its biological resources, with dire consequences. cal corollary of their dominant distribution in the
 northern hemisphere. The environmental services ures are implemented and maintained. The assess-
of natural conifer forests, coupled with the creation ment of conservation status of species on a global
of habitat to numerous species of animals, fungi and scale is largely undertaken by the International
plants and the provision of material and immaterial ­Union for the Conservation of Nature (IUCN). This
goods to human society are increasingly understood Non-Governmental Organisation (NGO), based in
as vital. The proper management of these forests Gland, Switzerland but operating worldwide, works
aims at the perpetuation of the original ecosystem on this through its Species Survival Commission
on large spatial and temporal scales and at least in (SSC) and numerous Specialist Groups, made up of
the temperate climate zones of the world utilization biologists and other specialists with special knowl-
is moving in the direction of such true sustainability. edge of certain groups of animals, plants, or other
30 However, recent Food and Agriculture Organisation defined groups. The Conifer Specialist Group is re-
of the United Nations (FAO) figures are showing sponsible for conifers in this context; it has a mem-
that subtropical and tropical coniferous forests are bership of around 40 individuals and is currently
declining faster than any other forest types. Outside chaired by the author of this Handbook. The con-
a limited number of reserves, exploitation not uti- servation status of a species involves knowledge of
lization is still the rule in much of the tropics. In- the extent of occurrence (EOO) in nature (exclud-
trinsic values to conifers may provide incentives to ing all introduced, planted and naturalized occur-
make sure they do not become extinct due to our rences) and the area of occupancy (AOO) within
activities. These can be aesthetic, cultural, emotional the natural range, both calculated in square kilom-
and scientific. People in all cultures show a remark- eters. These two factors determine how widespread
able veneration for individual trees, especially if (EOO) and how common (AOO) a species is. Popu-
these are old, large, or grow in special places. Tho- lation size is measured as numbers of mature (repro-
mas Pakenham, writer, photographer and planter ducing) individuals in total, or in subpopulations
of trees on his estate in Ireland, gives many exam- when occurring in scattered or fragmented groups
ples from many countries in his books on remark- of individuals. Counting individuals of trees is only
able trees, quite a number of which are conifers. The practicable if there are not too many, so many esti-
popularity of his books, illustrated with his similarly mates are based on EOO and AOO plus numbers of
remarkable photographs, amply demonstrates the subpopulations. The conservation status is also in-
fascination people still have with large and ancient fluenced by trends in the population: are the num-
individual trees. Many conifer species are apparent bers of individuals declining, stable or increasing?
relicts of a geological past when they, or their clos- Past, present, as well as projected future trends are
est relatives, were still abundant. When introduced of concern and can be used in a conservation assess-
by man to other continents, sometimes returning to ment. IUCN has developed criteria for conservation
where they once grew, some have done well. Perhaps assessment of this kind, which gives a species a rat-
therein lies the main reason to conserve them, it of- ing on a categorical scale of conservation status. An
fers the opportunity to restore something of a lost assessed and evaluated species is then added to the
world. There is also an obvious scientific interest, a IUCN Red List™ with its category of threat and the
curiosity value if you like, in these distinct species criteria used. The development of these criteria has
representing the past. Given the threat of extinction taken considerable time because a single system had
to so many species, prioritization of the species in to be made to work for all organisms. As a result,
need of conservation may be necessary to save at there have been two versions (dated 1994 and 2001)
least some of them. Phylogenetic distinction might which are currently both valid in terms of having
be a criterion that is well worth using to shortlist added species to the Red List. The Conifer Special-
species for conservation action. ist Group assessed most of the (now) 615 recognized
conifer species in the period between 1994 and 2001
The first task before us if we want to address the in- and only about 100 since then, some of the latter
creasing threats of extinction of conifer species is to a second time. These assessments are cited in this
assess the conservation status of these species. Types book. Re-assessment of the ca. 500 earlier assess-
of threat need to be identified, with their causes if ments is urgent, but was still in progress at the time
known. And finally, we would have to recommend of publication of this Handbook. The categories are
action and work to ensure these conservation meas- now as follows:
Extinct (EX) status. If these two causes are separated, it will be
Extinct in the Wild (EW) possible to see a trend in the overall conservation
Critically Endangered (CR) status of conifer diversity. This will then result in a
Endangered (EN) Red List Index for conifers, which will be a measure
Vulnerable (VU) of success, or failure, of conservation action because
Near Threatened (NT) it will tell us in what direction this diversity is mov-
Least Concern (LC) ing. It is very important that several non-related
Data Deficient (DD) taxonomic groups of organisms with worldwide dis-
Not Evaluated (NE) tributions, geographically aa well as ecologically, are
being assessed as completely as possible and as soon
The abbreviations are used with all species, subspe- as possible. The Convention on Biological Diversity 31
cies, and varieties treated in this Handbook. The cod- (CBD) has recommended to develop a number of
ed (abbreviated) criteria are also given, but their full indicators, including one based on changes in status
explanation would be too extensive to repeat here of threatened species, to be able to monitor progress
and the reader is referred to the originals (IUCN, towards the target of reducing the rate of loss of
1994, 2001). Taxa assessed and evaluated to be in any biological diversity (Baillie et al., 2004). IUCN has
of the three categories printed in bold are the taxa therefore developed Red List Indices for two such
threatened with extinction in a foreseeable future if groups: Birds and Amphibians. More taxonomic
trends of decline continue and if no adequate meas- groups with repeated complete assessments are ur-
ures are taken to alleviate the situation. As calculated gently needed in order to provide data for the in-
when writing this in April 2009, 230 of 794 accepted vestigation of trends in the conservation status of
taxa (species, subspecies and varieties) or 29% were biodiversity. The only taxonomic groups of plants
threatened with extinction. It is slightly more com- with suitable baseline data at present (2009) are the
plicated to calculate how many species would fall Cycads and the Conifers. Conifers are more valuable
in this broad category, because a species may have for this purpose than Cycads, of which the distribu-
one variety at risk and another variety not at risk. tion, although wide, is much more restricted both in
The categories of threat are therefore assigned to the terms of geography and of ecology. Conifers occur
lowest rank in case of a subdivided species. Only if indeed worldwide and occupy nearly all the major
all varieties recognized in a species are at risk, the biomes of the terrestrial world. It is hoped that the
entire species is at risk. This is true for ca. 180 or attention this Handbook may bring to the plight of
30% of all conifer species. These figures are likely to conifers in the world will help to put adequate and
change as species are re-assessed, partly due to more sufficient conservation policies and action in place
and better data, but also due to real changes in their to indeed slow down their extinction rate.
SYNOPSIS OF FAMILIES AND GENERA
This synopsis gives brief character sketches of taxa
Agathis Salisb.
in the ranks of genus to family. Only those char-
acters are mentioned which together would suffice Monoecious trees. Leaves subopposite to opposite,
to identify the taxon; these are not necessarily the leaf lamina broad and flat, with distinct petiole. Pol-
same for all taxa, and comparisons between them len cones solitary, relatively small. Seed cone scales 33
are therefore limited to those necessary for identi- with an imbricate, rounded margin. Seeds with a sin-
fication. In this Handbook 8 families and 70 gen- gle broad wing, becoming detached from the scale.
era are recognized within the order Coniferales, or
conifers. The families are here given in alphabetical
Wollemia W. G. Jones et al.
order, as no satisfactory classification at this level
based on all necessary evidence seems possible (Far- Monoecious trees. Leaves helically attached, sessile,
jon, 2007, 2008). The hypothetical relationships of adult leaves arranged in 4 rows (tetrastichous), linear.
genera within families are indicated by their group- Pollen and seed cones terminal on primary branch-
ings; some of these groupings have been named at es, relatively small. Seed cone scales with an apical
the rank of subfamily, others are here presented as free extension. Seeds circumferentially winged, re-
informal and separated from each other by a dou- maining attached to the scale; wing(s) narrow.
ble space only. The relationships in Podocarpaceae
are most tentative, as they are based on phylogenies
Araucaria Juss.
derived from recent cladistic analyses of molecular
and morphological data, which are not in agree- Dioecious or monoecious trees. Leaves in helical ar-
ment for several clades. The DNA-based analyses rangement, sessile, lamina broad and flat, or scale-
tend to place Phyllocladaceae within Podocarpaceae. like, persistent on falling branches. Pollen cones
Perhaps it could be assumed that the former arose solitary or in small clusters, relatively small to very
from the latter. The names of families and genera are large. Seed cone scales with an apical free extension.
given with their abbreviated authorities following Seeds without wings, remaining attached to the scale.
Brummitt & Powell (1992). For more detailed recent
classifications down to species see for Cupressaceae
Cephalotaxaceae Neger
Farjon (2005a), for Pinaceae excluding Pinus Farjon
(1990), for Pinus Richardson (ed., 1998) and Farjon
Cephalotaxus Siebold & Zucc. ex Endl.
(2005b) and for Taxaceae Cope (1998).
Dioecious evergreen shrubs or small trees. Leaves
pectinately arranged becoming subopposite, linear-
Araucariaceae Henkel & W. Hochst.
lanceolate, with two prominent stomatal bands on
Dioecious or monoecious evergreen, highly resin- the abaxial side. Pollen cones aggregated in capitu-
ous trees. Tree architecture according to Massart’s lae. Seed cones reduced, with opposite fertile bracts.
and Rauh’s models. Leaves in helical arrangement or Seeds 1–2 per cone, exposed, large, completely sur-
subopposite to opposite; lamina broad and flat, or rounded by a fleshy aril.
scale-like. Pollen cones catkin-like, sometimes large.
Seed cones large, globose, mostly disintegrating.
Cupressaceae Gray
Seed cone scales predominantly consisting of the
bract, but with a fused seed scale bearing a single Aromatic, evergreen or deciduous, monoecious or
seed. dioecious shrubs or trees. Leaves on (pen)ultimate
branchlets linear, needle- or scale-like, spirally ar-
ranged, ternate or decussate (rarely quadrate) in
mature plants. Pollen cones small, terminal, rarely
 axillary, solitary or sometimes clustered in groups of
Metasequoia Hu & W. C. Cheng
2–7. Seed cones terminal, simple or semi-compound,
globose, ovoid or conical. Seed cone scales consist- Deciduous, monoecious trees, dropping foliage
ing of transformed bracts, true seed scales absent or branchlets, not individual leaves. Leaves opposite,
sometimes rudimentary. Seeds 1-many axillary to linear, spreading at nearly right angles to the shoot.
each cone scale, with or without wings. Pollen cones numerous in spike-like shoot systems.
Seed cones terminal on 2–5 cm long, scale-leaved
shoots, subglobose, barrel-shaped or fusiform.
Cunninghamioideae (Zucc. ex Endl.) Quinn
Seeds numerous, with 2 marginal wings.
Cunninghamia R. Br.
34
Sequoia Endl.
Evergreen, monoecious trees, capacity to coppice
profound. Leaves helically arranged, linear-lanceo- Evergreen, monoecious (very tall) trees, often
late; leaf margin serrulate. Pollen cones numerous in sprouting from lignotubers. Leaves alternate, most-
clusters. Seed cones subterminal, persistent (falling ly linear, pectinate on shaded shoots. Pollen cones
with foliage branches), with thin, coriaceous scales. on the same branches as seed cones, solitary. Seed
Seeds 2–3 per fertile scale, with 2 marginal wings cones terminal on short branchlets, more or less
1 mm wide. ovoid, 15–30 mm long. Bract-scale complexes heli-
cally arranged, parting to release the marginally
winged seeds.
Taiwanioideae L. C. Li

Taiwania Hayata Sequoiadendron J. Buchholz

Evergreen, monoecious trees. Leaves helically ar- Evergreen, monoecious (giant) trees. Leaves heli-
ranged, imbricate, in young trees falcate-subulate, cally arranged in 3 ranks, imbricate, scale-like. Pol-
ultimately scale-like. Pollen cones in terminal clus- len cones on the same branches as seed cones but
ters on branchlets with scale leaves. Seed cones ter- well above them, solitary. Seed cones terminal on
minal, solitary, small, with thin, coriaceous scales. short branchlets, 30–70(–95) cm long. Bract-scale
Seeds usually 2 per fertile scale, each with 2 small complexes helically arranged, parting to release the
wings. flattened, unequally winged seeds.

Athrotaxoideae L. C. Li Taxodioideae Endl. ex K. Koch

Deciduous or evergreen, monoecious trees. Leaves
Athrotaxis D. Don
alternate to helically arranged, of various types but
Evergreen, monoecious trees. Leaves helically ar- linear leaves present. Pollen cones crowded or ar-
ranged, small and appressed or longer than 6 mm ranged in spike-like to paniculate systems, solitary.
and spreading gradually or abruptly. Pollen cones Seed cones terminal, solitary or clustered; bract-
solitary, small. Seed cones terminal, solitary, globose scale complexes with or without small teeth below
or subglobose when opened, with clavate-peltate, the apex. Seeds with 1–2 wings, or almost wingless.
thin or thick woody scales. Seeds with two slightly
unequal wings.
Cryptomeria D. Don
Evergreen, monoecious trees. Leaves helically ar-
Sequoioideae Saxton
ranged in 5 ranks, decurrent, free for 1/2–3/4 of leaf
Deciduous or evergreen, monoecious trees. Leaves length, linear-subulate. Pollen cones numerous, ax-
helically arranged or opposite, linear or scale-like. illary and crowded towards the ends of branchlets.
Pollen cones solitary or numerous in spike-like Seed cones terminal, often aggregated; bract-scale
shoot systems. Seed cones barrel-shaped or ovoid, complexes helically arranged, spreading, with a
with peltate bract-scale complexes. Seeds axillary to number of small teeth below the bract apex. Seeds
cone scales, with 2 marginal wings. with 2 narrow, unequal wings.
 Seed cones ovoid-globose, narrowly ovoid or ellip-
Glyptostrobus Endl.
tical, partly opening; bract-scale complexes decus-
Semi-evergreen, monoecious trees. Leaves alternate sate, the lowest pair reduced, the 2–3 following pairs
to helically arranged, consisting of 3 types, scale-like, enlarged and spreading, the apical pair often con-
subulate, and linear. Pollen cones terminal, solitary. nate. Seeds with 2 wings.
Seed cones terminal, solitary, pyriform to obovate;
bract-scale complexes obovate-oblong, connate but
Calocedrus Kurz
parting slightly at maturity, with many tiny, sub­
apical tooth-like lobes. Seeds with a single wing. Evergreen, monoecious trees. Leaves decussate, ap-
pearing in whorls of 4, imbricate, scale-like, dimor-
phic, decurrent. Pollen cones solitary, cylindrical. 35
Taxodium Rich.
Seed cones solitary; bract-scale complexes in 3(–4)
Deciduous or semi-deciduous, monoecious trees, decussate pairs. Seeds 1–2 per fertile scale, with 2
dropping foliage branchlets, not individual leaves. very unequal wings.
Leaves helically arranged, spreading in 2 ranks or
in 5–8 ranks, linear or (short) acicular. Pollen cones
Chamaecyparis Spach
arranged in spike-like to paniculate systems. Seed
cones terminal and often clustered, subglobose or Evergreen, monoecious trees. Leaves decussate,
more or less ovoid; bract-scale complexes breaking imbricate, scale-like, dimorphic. Pollen cones soli-
away to release the almost wingless seeds. tary, short cylindrical, with yellow or red pollen
sacs. Seed cones solitary, globose to ellipsoid-ovoid,
small, with ± peltate, parting scales. Seeds with 2 lat-
Cupressoideae Rich. ex Sweet
eral, narrow wings.
Evergreen, dioecious or monoecious trees. Adult
leaves scale-like or acicular; scale leaves dimorphic
Cupressus L.
or monomorphic mostly dependent on plagiotropic
or orthotropic branching, often with a gland. Pol- Evergreen, monoecious shrubs or trees. Leaves
len cones terminal or sometimes axillary, very small, decussate, scale-like, monomorphic or dimorphic.
solitary or sometimes 2–3 together. Seed cones ter- Pollen cones solitary, short cylindrical, with yellow
minal on foliage branchlets or on dwarf shoots, pollen sacs. Seed cones often grouped close together
variable in size and shape; bract-scale complexes or clustered, globose to ovoid-oblong with parting
3–4 whorled or decussate, valvate, peltate, or fused. woody scales, caducous or persistent; bract-scale
Seeds 1-many per cone, winged or wingless. complexes decussate, peltate. Seeds numerous, an-
gular, with rudimentary wings.
Thujopsis Siebold & Zucc. ex Endl.
Fokienia A. Henry & H. H. Thomas
Evergreen monoecious trees. Leaves decussate,
imbricate, scale-like, dimorphic at least on plagio- Evergreen, monoecious trees. Leaves decussate, im-
tropic branchlets, lustrous green, with broad, white bricate, scale-like, (strongly) dimorphic, decurrent.
stomatal bands on the underside of branchlets. Pollen cones solitary, subglobose to oblong. Seed
Pollen cones ovoid-globose, reddish purple. Seed cones subglobose; bract-scale complexes decussate,
cones subglobose, markedly umbonate, 8–16(–20) spreading at maturity. Seeds with 2 very unequal
mm long; bract-scale complexes decussate, apically wings.
thickened. Seeds with 2 wings.
Juniperus L.
Thuja L.
Evergreen, monoecious or dioecious shrubs or
Evergreen, monoecious shrubs or trees. Leaves dec- trees. Leaves decussate or ternate, scale-like or ac-
ussate, imbricate, scale-like, dimorphic on lateral icular, decurrent or articulate. Pollen cones terminal
branchlets. Pollen cones oblong, yellowish green. or axillary. Seed cones terminal on axillary (dwarf)
 shoots, globose, semiglobose or ovoid, indehiscent, leaves, ovoid-oblong; bract-scale complexes 4, dec-
usually red, glaucous or blue; bract-scale complexes ussate, the upper scales twice as large as the lower.
in 1–4 decussate or ternate whorls, entirely fused, Seeds with 2 wings, one rudimentary, the other well
usually soft. Seeds 1–8 per cone, wingless. developed.

Microbiota Kom. Libocedrus Endl.

Evergreen, monoecious, decumbent shrubs. Leaves Evergreen, monoecious shrubs or trees. Leaves
decussate, imbricate, scale-like, weakly dimorphic, decussate, imbricate, scale-like, decurrent, strong-
turning copper-brown or purplish brown in winter. ly dimorphic. Seed cones terminal on flattened or
36 Pollen cones axillary. Seed cones hidden in shoot quadrangular branchlets, subtended by 4–5 decus-
axils, minute, reduced to 2 pairs of bract-scales, the sate, transitional leaf pairs; bract-scale complexes
upper enclosing a single, ovoid, wingless seed. in 2 decussate pairs, the upper fertile pair of scales
spreading at maturity, with a long subapical bract
tip. Seeds 1–4 per cone, with 2 very unequal wings.
Platycladus Spach
Evergreen, monoecious trees. Leaves decussate,
Papuacedrus H. L. Li
scale-like, decurrent, on lateral branchlets dimor-
phic. Pollen cones solitary, subglobose. Seed cones Evergreen, monoecious trees, often appearing dioe-
erect, ampulliform, opening widely; bract-scale cious. Leaves in whorls of 4 or decussate, on lateral
complexes 6–8, decussate. Seeds ovoid; wings absent branchlets scale-like, strongly dimorphic. Pollen
or rudimentary. cones solitary, cylindrical. Seed cones (sub)termi-
nal, thin woody; bract-scale complexes consisting of
2 decussate pairs, the lower curved, the upper much
Tetraclinis Mast.
larger, elliptic. Seeds 2–4 per cone, with 2 very un-
Evergreen, monoecious shrubs or small trees. equal wings.
Leaves decussate, scale-like, long decurrent, adnate
to shoot except the apex, weakly dimorphic. Pollen
Pilgerodendron Florin
cones solitary, ovoid-globose, reddish. Seed cones
(sub)tetragonal; bract-scale complexes 4, decussate, Evergreen, dioecious trees. Leaves decussate, occa-
nearly equal in size, thick woody. Seeds with 2 large, sionally in whorls of 3, imbricate, forming 4(6) rows,
thin membranous wings. scale-like, monomorphic. Seed cones terminal, sub-
tended by 2 decussate, transitional leaf pairs, with
distally spreading scales; bract-scale complexes in 2
Xanthocyparis Farjon & Hiep
decussate pairs, elliptic to obtrullate, the upper pair
Evergreen, monoecious trees. Leaves scale-like twice as large as the lower pair; columella conspicu-
(adult) and acicular-linear (juvenile), in one species ous. Seeds 3–4 per cone, with 2 very unequal wings.
both occur in mature trees; adult leaves decussate,
dimorphic, juvenile leaves in whorls of 4, mono-
Actinostrobus Miq.
morphic. Pollen cones solitary, yellow. Seed cones
opening wide; bract-scale complexes 4 (or 6) in dec- Evergreen, monoecious shrubs or trees. Leaves in
ussate pairs. Seeds flattened, with 2 lateral wings. whorls of 3, decurrent; juvenile acicular leaves on
young plants only or also on mature plants; adult
leaves mostly shorter than 5 mm, linear-lanceolate.
Seed cones persistent along branches and stems;
Austrocedrus Florin & Boutelje
bract-scale complexes in two whorls of 3, of nearly
Evergreen, dioecious shrubs or trees. Leaves decus- equal size at maturity, subtended by 4–6 alternating
sate, in opposite ranks, scale-like, dimorphic; facial whorls of 3 broad, imbricate scale leaves; columella a
leaves very small. Pollen cones solitary, small. Seed strong, acute spike. Seeds with 3 wings.
cones on ultimate branchlets with non-modified
 complexes 4 (rarely 6), valvate, in two pairs of slight-
Callitris Vent.
ly unequal size and shape, thick woody; columella
Evergreen, monoecious shrubs or trees. Leaves in short, sometimes double. Seeds numerous, with or
alternate whorls of 3 (in one species also with ac- without 2 wings.
icular, spreading leaves in whorls of 4), scale-like,
­decurrent, appressed, linear. Pollen cones solitary
Phyllocladaceae Bessey
or with 2–3 together at the tips of branchlets. Seed
cones solitary, opening soon and deciduous, but
Phyllocladus Rich. ex Mirb.
more often clustered and persistent and often se-
rotinous; bract-scale complexes in two alternate Evergreen shrubs or trees, usually dioecious, oc-
whorls of 3 (or 4), the upper whorl usually the larg- casionally monoecious. Lateral branches of highest 37
est; columella distinct, often trimerous. Seeds with order transformed to simple or compound (pinnate)
2–3 wings. phylloclades functioning as leaves, arranged spirally
or in pseudo-whorls. True leaves rudimentary. Seed
cones on petiolate or foliate parts of phylloclades,
Diselma Hook. f.
arranged in rows, in pairs or solitary. Seeds ovoid,
Evergreen, dioecious shrubs, or very small trees. partly embedded by the fused, swollen and succu-
Leaves opposite-decussate, occasionally in whorls of lent bract scales of the cone and subtended by a filmy
3, scale-like, imbricate. Pollen cones numerous, soli- white aril leaving the apical part of the seed free.
tary. Seed cones terminal, maturing in one season to
small cones of 3 × 2 mm, consisting of 2 pairs of dec-
Pinaceae Spreng. ex F. Rudolphi
ussate cone scales surrounding a central columella.
Seeds 2 (or 1), with 2(–3) marginal wings. Monoecious evergreen or deciduous, resinous trees
or shrubs. Leaves spirally inserted on long or short
shoots and solitary or arranged in fascicles of 1–8
Fitzroya Lindl.
surrounded by a sheath on dwarf shoots (Pinus),
Evergreen, dioecious trees. Leaves in alternate acicular-linear to long linear. Pollen cones often
near-whorls of 3, imbricate, scale-like, decurrent. grouped close together on long shoots, axillary, soli-
Seed cones formed by 2–3 whorls of slightly modi- tary or clustered from a single bud, catkin-like, de-
fied scale leaves, followed by 2 alternate whorls of ciduous. Seed cones compound, small to very large.
3 fertile, wide spreading scales; columella variably Bracts free, well developed or rudimentary. Seed
shaped, trigonal to tripartite. Seeds with 2–3 narrow scales persistent or deciduous, woody; bearing 2 in-
wings. verted seeds. Seeds with 1 wing or wingless.

Neocallitropsis Florin Pinoideae Pilg.

Evergreen shrubs or small trees (distribution of sex- Evergreen trees or shrubs. Leaves spirally inserted
es uncertain). Leaves in alternate whorls of 4, seem- on long shoots or in fascicles of 1–8 on dwarf shoots
ingly in 8 rows, short decurrent or nearly sessile with (Pinus), acicular or (long) linear. Pollen cones
broad base, lanceolate, incurved, thick. Pollen cones grouped close together on long shoots (Pinus) or
10–12 × 6–7 mm, subglobose to ovoid. Seed cones solitary in leaf axils. Bracts of seed cones small but
with spreading bract-scale complexes; columella conspicuous or rudimentary. Seed scales persist-
short pyramidal. Seeds with 2(–3) marginal wings. ent, thin or thick woody. Seeds winged or wingless.
Number of cotyledons in seedlings high (4-≥20).
Widdringtonia Endl.
Pinus L.
Evergreen, monoecious shrubs or trees. Leaves dec-
ussate on smallest branchlets, becoming more or Trees or shrubs. Leaves in fascicles of 1–8 (com-
less spirally arranged (bijugate) on thicker branch- monly 2, 3 or 5) on dwarf shoots, long linear. Pollen
lets, scale-like. Pollen cones solitary. Seed cones cones grouped close together in helical arrangement
usually clustered (to 50 mature cones); bract-scale on long shoots. Seed cones at base of new shoots,
 small to very large, persistent or deciduous. Bracts ils of leaves. Seed cones becoming pendant at matu-
rudimentary. Seed scales usually thick woody, rity, falling entire from branches with the peduncle
sometimes remaining soft, with a distinct apophysis attached. Bracts exserted, longer than seed scales,
and umbo. Seeds winged or wingless, wing adnate with trilobate apex. Seed scales with a rounded up-
or easily detached. per margin. Seeds winged; wing short.

Cathaya Chun & Kuang Abietoideae Pilg.

Trees. Leaves on short shoots seemingly in tufts, on Evergreen or deciduous trees. Leaves on long shoots
long shoots more remote, linear, flattened, longitu- and/or on short shoots, linear or acicular. Pollen
38 dinally grooved above. Pollen cones lateral, near the cones solitary or clustered, in axils of leaves or ter-
shoot apex, pendant. Seed cones lateral, ovoid ob- minal on short shoots. Seed cones erect and disin-
long, with acute apex, 3–5 cm long. Bracts triangu- tegrating when fully mature, or becoming pendant
lar, with a long point. Seed scales 13–16, thin woody, and falling intact. Bracts well developed, small, or
orbicular or ovate. Seeds ovoid, winged; wing 2 × rudimentary. Seed scales with narrowed, pedicellate
length of seed, adnate. base. Seeds winged, with resin vesicles in seed coat.

Picea A. Dietr. Abies Mill.

Trees. Leaves inserted (adnate) on pulvini on long Evergreen trees. Leaves on long shoots, linear, leav-
shoots, acicular. Pollen cones lateral, numerous but ing a round scar when falling. Pollen cones solitary
solitary in leaf axils. Seed cones becoming pendant, in axils of leaves of ultimate branchlets. Seed cones
deciduous. Bracts rudimentary. Seed scales numer- erect, yellowish green to deep purple or dark blue,
ous, thin woody. Seeds winged; wing usually 3 × often exuding resin. Bracts exserted or included.
length of seed, easily detached. Seed scales falling with the broadly winged seeds,
leaving the erect rachis on the tree.
Laricoideae Melchior & Werdermann
Cedrus Trew*
Evergreen or deciduous trees. Leaves on long shoots
and/or short shoots, linear. Pollen cones lateral in Evergreen trees with pronounced shoot dimor-
leaf axils, or terminal and clustered on short shoots. phism. Leaves on long shoots but predominantly
Seed cones pendant or erect, deciduous or falling at- on short shoots, acicular. Pollen cones solitary at
tached to branches. Bracts well developed, often ex- the apex of short shoots, with anthesis in autumn.
serted. Seed scales thin woody. Seeds winged; wing Seed cones remaining erect, maturing in second
2 × length of seed, adnate. year. Bracts rudimentary. Seed scales falling with
the broadly winged seeds, leaving the erect rachis on
the tree.
Larix Mill.
Deciduous trees with pronounced shoot dimor- * The genus Cedrus has come out as basal (sister
phism. Leaves on long shoots but predominantly on group) to all other genera of the Pinaceae in some
short shoots, linear, lax. Pollen cones terminal and recent DNA-based cladistic analyses; morphologi-
clustered on short shoots. Seed cones remaining cal evidence however places it firmly within Abieto­
erect, persistent and separated from the tree while ideae. A basal position is not corroborated by the
still attached to short shoots and branches. Bracts fossil record. Its position is tentative.
exserted or included, not or only slightly longer than
seed scales.
Keteleeria Carrière
Evergreen trees. Leaves on long shoots, broadly lin-
Pseudotsuga Carrière
ear to lanceolate-linear. Pollen cones solitary in leaf
Evergreen trees. Leaves on long shoots, leaving axils. Seed cones remaining erect, breaking up by
small scars when falling. Pollen cones solitary in ax- disintegration of the rachis. Bracts well developed,
often exserted. Seed scales persistent. Seeds with
Saxegothaea Lindl.
broad wing dispersed separately; seed germination
hypogeal. Monoecious trees. Leaves pectinate, short decur-
rent, linear or falcate. Pollen cones axillary, solitary
or sometimes in pairs, 4–6 mm long. Seed cones ter-
Nothotsuga Hu ex C. N. Page
minal, globular, 9–12 mm diam. Cone scales 15–20
Evergreen trees with weakly developed shoot di- per cone, spirally arranged, swelling at maturity.
morphism. Leaves linear. Pollen cones in umbellate Seeds 1 per scale (from 2 inverted ovules), enclosed.
clusters from a single bud. Seed cones remaining
more or less erect, falling entire or disintegrating.
Podocarpus L’Hér. ex Pers.
Seed scales persistent. Seeds with small wing dis- 39
persed separately. Dioecious or rarely monoecious shrubs or more
commonly trees. Leaves relatively broad, usually
linear-lanceolate or linear-elliptic, coriaceous, with
Pseudolarix Gordon
a single raised, flat or sunken midrib and stomata in
Deciduous trees with pronounced shoot dimor- two broad bands on the abaxial side. Pollen cones
phism. Leaves on long shoots but predominantly on axillary, solitary or clustered, catkin-like. Seed cones
short shoots, linear, lax. Pollen cones in umbellate swelling to form a smooth, succulent, coloured re-
clusters from the apex of a short shoot. Seed cones ceptacle. Seeds 1(–2) per cone, completely exposed,
terminal on short shoots, remaining erect, disin- drupe-like, completely covered by a fleshy epima-
tegrating. Seed scales persistent, narrowly deltoid- tium.
triangular, subtended by a small bract. Seeds with a
large, narrow wing.
Retrophyllum C. N. Page
Dioecious, dwarfed to large trees. Leaves spirally
Tsuga (Endl.) Carrière
inserted, lanceolate to narrowly ovate, obliquely
Evergreen trees with weakly developed shoot dimor- directed into subopposite and decussate apparent
phism. Leaves on long shoots (some rather small), pairs by twisted petioles in opposite directions, am-
linear, with a bent and twisted petiole. Pollen cones phistomatic. Pollen cones axillary or terminal. Seed
axillary, solitary, small. Seed cones pendulous, fall- cones rudimentary, with a single large seed covered
ing entire without branchlets or peduncles attached. by a fleshy, drupe-like epimatium.
Seed scales thin, with rounded margin. Seeds with a
relatively large, oblique wing.
Afrocarpus (J. Buchholz & N. E. Gray)
C. N. Page
Podocarpaceae Endl.
Dioecious trees. Leaves twisted in opposite direc-
Dioecious or sometimes monoecious, evergreen tions, narrowly lanceolate-elliptic to linear-lanceo-
trees or (dwarf) shrubs (one species parasitic on late, coriaceous, with a single midrib, amphistomatic.
another member of the family). Leaves helically ar- Pollen cones axillary, solitary or in groups of 2–3,
ranged, rarely decussate (Microcachrys), appressed catkin-like. Seed cones much reduced, not trans-
and imbricate to spreading and remote, shapes formed into a receptacle. Seeds 1 per cone, subtend-
highly variable, ranging from small, appressed scale ed by small, withering scales, entirely enclosed by a
leaves, via thin acicular leaves, to dorsiventrally flat- swollen, drupe-like epimatium.
tened, linear-lanceolate to broadly lanceolate, large
leaves (up to 34 × 9.5 cm). Pollen cones mostly sim-
Nageia Gaertn.
ple, axillary or terminal, solitary or clustered. Seed
cones axillary or terminal, solitary, mostly much Dioecious or monoecious trees, rarely shrubs.
reduced and often swelling to form a succulent re- Leaves large, flat, broadly ovate-elliptic to lanceolate,
ceptacle; fertile bracts 1-many. Seeds of most species lacking a midrib and with many parallel, converging
surrounded by a coriaceous or succulent epimatium. veins. Pollen cones single or in small, spicate groups
 of 2–6 on axillary peduncles, ovoid-cylindric. Seed
Halocarpus Quinn
cones much reduced, or forming a weakly devel-
oped, fleshy receptacle. Seeds 1 per cone, exposed, Dioecious shrubs or trees. Leaves dimorphic, radi-
entirely enclosed by a swollen, drupe-like epima- ally spreading, lanceolate-linear to linear ones alter-
tium. nating with appressed, rhombic scale leaves. Pollen
cones solitary or with 2–3 together at the apex of
scale-leaved branchlets, small. Seed cones reduced
to a few reddish bracts at the apex of scale-leaved
Acmopyle Pilg.
branchlets. Seeds 1–5 per cone, erect, at base sur-
Dioecious (or sometimes monoecious?) small trees. rounded by a swollen epimatium forming a white or
40 Leaves dimorphic, small and scale-like on leading yellow collar, lustrous black.
and fertile shoots, larger and falcate-linear on later-
al, vegetative shoots. Pollen cones cylindrical, more
Lagarostrobos Quinn
or less erect. Seed cones solitary or occasionally
grouped, when mature forming an irregular, fleshy Predominantly dioecious trees, with lower branches
and verrucose receptacle. Seeds single, enclosed by frequently layering. Adult leaves imbricate and ap-
a fleshy epimatium. pressed, rhomboid in appearance, 1–1.5 × 1 mm. Pol-
len cones terminal, sessile. Seed cones terminal on
decurved short branchlets, 4–5 mm long. Seeds up
Dacrycarpus (J. J. Bennett) de Laub.
to 5–8 per cone, usually fewer, light brown, enclosed
Dioecious or rarely monoecious shrubs or trees. at base by a dry, papery epimatium.
Leaves trimorphic, with small scale leaves, acicu-
lar leaves and flattened, linear-falcate leaves. Pol-
Lepidothamnus Phil.
len cones single or in pairs on axillary short shoots,
­cylindrical. Seed cones solitary, when mature form- Dioecious or sometimes monoecious, creeping or
ing an irregular, fleshy and verrucose receptacle. erect shrubs or trees. Leaves dimorphic, with lin-
Seeds single, enclosed by a fleshy epimatium. ear, spreading juvenile ones usually gradually giving
way to appressed, ovate-rhombic and gibbous scale
leaves. Pollen cones terminal or sometimes lateral.
Dacrydium Sol. ex G. Forst.
Seed cones terminal, solitary, very small, consist-
Dioecious or rarely monoecious shrubs or trees. ing of 3–5 yellow or reddish bracts. Seeds 1 per cone,
Leaves dimorphic, scale-like and subulate or acicu- purplish brown or black, surrounded at base by a
lar. Pollen cones small, cylindrical. Seed cones small, membranous epimatium.
forming a red receptacle. Seeds 1–2(–3) per cone, be-
coming more or less erect, protruding from a cup-
Manoao Molloy
like epimatium covering basal part of seed only,
lustrous brown or nearly black. Predominantly dioecious trees; ‘sucker’ shoots from
horizontal underground stems prolific. Adult leaves
imbricate and appressed, rhomboid in appearance,
Falcatifolium de Laub.
1–1.5 × 1 mm. Pollen cones terminal, sessile. Seed
Dioecious shrubs or trees. Leaves dimorphic, with cones terminal on distally curved short branchlets,
scale leaves on leading and fertile shoots, alternating 3–4 mm long. Seeds 1–3(–5) per cone, dark purple to
with more or less distichously spreading, bilaterally black, the basal half or more enclosed by a swollen,
flattened, obliquely lanceolate-falcate leaves on veg- fleshy epimatium.
etative shoots. Pollen cones cylindrical and catkin-
like. Seed cones forming an irregular, red receptacle.
Microcachrys Hook. f.
Seeds erect, surrounded at the base by a swollen epi-
matium, with two lateral ridges. Monoecious or (temporarily) dioecious, prostrate
shrubs. Leaves on lateral branchlets decussate, im-
bricate, appressed, short triangular in appearance,
1–1.5 × 1 mm. Pollen cones terminal, more or less leaves much reduced, scale-like. Pollen cones in ter-
recurved. Seed cones terminal, with ca. 20 helically minal or sublateral clusters. Seed cones terminal or
arranged, fleshy, red bracts. Seeds 1 per fertile bract sublateral, ovoid to cylindrical, often exuding resin.
scale, at base partly covered by an epimatium. Bract-scale complexes helically arranged, consist-
ing of a partly fused bract and exceeding seed scale.
Seeds basal to seed scales, inverted, with 2 wings.
Parasitaxus de Laub.
Small erect monoecious shrubs, parasitic on the
Taxaceae Gray
roots of Falcatifolium taxoides. Leaves spirally ar-
ranged, imbricate, scale-like, decurrent, reddish. Dioecious or rarely monoecious evergreen shrubs or
Pollen cones usually on the same branching systems trees. Leaves helically or decussately inserted, usu- 41
as seed cones, terminal, solitary. Seed cones reduced ally pectinately arranged, linear to lanceolate with
to a few bracts with a single seed surrounded by a a single midrib, hypostomatic, with two bands of
hard, glaucous white epimatium. stomata. Pollen cones axillary to foliage leaves and
solitary or aggregated in umbellate clusters of ra-
cemes. Seed bearing structures consisting of axillary
Pherosphaera W. Archer
dwarf shoots with terminal, erect ovules. Seeds sur-
Dioecious shrubs. Adult and juvenile leaves simi- rounded by a fleshy or succulent aril; aril partially or
lar, spirally arranged, 2–4(–6) mm long and mostly entirely enclosing the seed, becoming red, purple or
scale-like. Pollen cones terminal, globose to ovoid. yellow when ripe.
Seed cones terminal, with (3–)5–8 spreading, fertile
scales, not fleshy at maturity. Seeds usually 1–4 per
Amentotaxus Pilg.
cone, exposed, erect.
Dioecious shrubs or small trees. Leaves usually
opposite-decussate, pectinately arranged by twisted
petiolate leaf bases, linear-lanceolate, straight or
Prumnopitys Phil.
slightly falcate, large. Pollen cones aggregated in
Dioecious trees or shrubs. Leaves small, in lateral umbellate clusters of racemes. Seed-bearing struc-
shoots pectinately arranged, flattened, falcate-linear. tures usually in groups near the apex of vegetative
Pollen cones aggregate on relatively long, leafless shoots. Aril surrounding the ripe seed greatly en-
branchlets, cylindrical. Seed cones reduced to a few larged, ellipsoid or ovoid, fleshy and succulent, red
bracts. Seeds enclosed in a drupe-like, fleshy, yellow, or purple, completely hiding the small seed.
red or black epimatium.

Sundacarpus (J. Buchholz & N. E. Gray) Austrotaxus R. H. Compton

C. N. Page
Dioecious trees. Leaves alternate (helical), spread-
Dioecious trees. Adult leaves broadly linear, 5–15 cm ing forward at 30–60° to the shoot axis, narrowly
long, 6–15 mm wide, petiolate, with a single mid- lanceolate, large. Pollen cones axillary, solitary. Seed
vein. Pollen cones axillary, often 3–7 on a stalk, cy- aril almost entirely enclosing the seed and only leav-
lindrical. Seed cones much reduced, not forming a ing the seed apex free, ovoid, smooth, at first glau-
receptacle. Seeds 1 per cone, large, covered by a glau- cous green ripening to orange.
cous purple, fleshy but firm epimatium.

Sciadopityaceae Luerss. Pseudotaxus W. C. Cheng

Dioecious shrubs. Leaves helically arranged, dis-
Sciadopitys Siebold & Zucc.
tichous, spreading, linear. Pollen cones axillary,
Evergreen, monoecious trees. Foliage consisting solitary. Seed aril cupular, only partly enclosing the
of linear cladodes (‘needles’) in pseudo-whorls on seed, succulent and white when ripe.
shortened terminal sections of long shoots. True
 Taxus L.
Dioecious shrubs or trees. Leaves helically inserted,
twisted at petiolate base and becoming pectinate or
distichous, linear or falcate. Pollen cones axillary,
solitary. Seed aril cupular, only partly enclosing the
seed, succulent and red, orange or yellow when ripe.

Torreya Arn.
42
Dioecious or rarely monoecious trees or some-
times shrubs. Leaves helically inserted, pectinately
arranged, linear-lanceolate, rigidly coriaceous,
­acuminate or pungent. Pollen cones axillary, soli-
tary. Aril almost completely surrounding the large
seed, fleshy, purplish green to bluish black when
ripe.
TAXONOMIC TREATMENT OF FAMILIES

base of each bract (rarely only a single seed per

Key to the families
cone) 4
This Handbook recognizes eight families within the 3b. Seed cones either with much reduced, obscure
conifers (currently with 70 genera and 614 species); scales (whether bracts or seed scales, in a few
some of their diagnostic characters are here used in instances with many very small bracts) or lack- 43
the key to families. If the family to which a species ing any kind of scales and reduced to a single,
has been assigned is not known to the user, it is nec- terminal seed with a surrounding epimatium
essary to start with this key and then proceed to the or arillus 5
key to genera under the family determined. If the 4a. True leaves scale-like or acicular (needle-like),
family is known, one can proceed directly to the lat- cataphylls absent, phyllodes absent
ter key to determine the genus. Keys to species with- Cupressaceae
in genera are provided under each genus; the species 4b. True leaves reduced to tiny cataphylls (usually
are arranged in alphabetical order (A-Z) throughout brown scales); phyllodes (pseudo-leaves) in the
using the Latin binomials, which enables the user to axils of these green, needle-like, growing rhyth-
find the genus determined using the keys with ease. mically in pseudo-whorls on shoots
Sciadopityaceae
1a. Seed cones with seed scales in the axils of bracts 5a. Seed cones reduced to a single seed; seed ter-
(bracts can be much reduced in mature, woody minally placed at a scaly dwarf shoot, partly or
cones but are conspicuous in immature cones completely surrounded by a succulent or fleshy
at early stages of development); i.e. cones clear- aril Taxaceae
ly compound and never reduced; seeds two on 5b. Seed cones apparent, with much reduced,
the adaxial (upper) side of each fertile scale; obscure scales (usually bracts; if these are
adult green leaves acicular-linear Pinaceae seemingly absent, seed on a stout or fleshy ‘pe-
1b. Seed cones with seed scales fused with bracts duncle’) or in a few instances with many, very
(bracts make up the bulk of the cone), or with small bracts 6
bracts only (which may be much enlarged, 6a. Pollen cones aggregated in more or less globose
swollen and/or woody at maturity), or with capitulae (‘heads’); bracts of (reduced) seed
scales obscure, much reduced or absent; seeds cones decussate Cephalotaxaceae
either single or more than two per fertile scale; 6b. Pollen cones solitary or clustered but not in glo-
adult green leaves scale-like, or acicular, or with bose capitulae; bracts of (reduced) seed cones
a distinct lamina, or replaced by phylloclades alternate or helically arranged (but sometimes
(phyllodes) 2 seemingly absent being enveloped by a swollen
2a. Seed cones with seed scales fused with bracts, receptacle) 7
or with bracts only which are much enlarged 7a. True leaves scale-like, acicular or with a dis-
and often swollen at maturity and may then tinct lamina, always simple; cataphylls absent;
form a compact, globose cone 3 phylloclades absent; cones arising in the axils
2b Seed cones with seed scales fused with bracts; of leaves Podocarpaceae
bracts forming the largest part of the cone 7b. True leaves reduced to tiny cataphylls, usually
scales; a single inverted seed per scale soon deciduous; phylloclades (pseudo-leaves)
Araucariaceae leaf-like, green, simple or compound; cones
3a. Seed cones consisting of bracts forming the arising on the edges of phylloclades or on sepa-
cone scales only (sometimes with rudimentary rate determinate shoots (axillary to bracts)
seed scales only visible at very early stages of Phyllocladaceae
development); seeds 1-many, axillary or on the
 Descriptions of families and keys to genera
Three genera: Agathis (17 spp.), Araucaria (19 spp.)
and Wollemia (1 sp.); total 37 species.
Araucariaceae Henkel & W. Hochst., Syn.
Nadelhölz.: xvii, 1. 1865. (nom. cons.). Type:
Distribution
Araucaria Juss.
Malesia: all major islands except Jawa and Lesser
Sunda Islands; Australia: New South Wales, Queens-
Description
land. SW Pacific: New Caledonia, Vanuatu, Fiji,
Dioecious or monoecious evergreen, highly resin- Norfolk Island, New Zealand (North Island). South
44 ous trees. Tree architecture according to Massart’s America: SE Brazil, NE Argentina; S Chile, SW Ar-
and Rauh’s models. Resin canals in bark, leaves and gentina (Andes).
seed cones. Bark hard and smooth, exfoliating with
rounded or irregular flakes (Agathis), rough and ex-
Key to the genera
foliating in horizontal strips and eventually deeply
fissured (Araucaria), or forming many irregular 1a. Bark hard and smooth, exfoliating with round-
pustules (Wollemia). Branches in rhytmic pseudo­ ed or irregular flakes; leaves distinctly petiolate;
whorls, spreading and plagiotropic (Massart’s seed cones globose, with imbricate scales lack-
model) or ascending to become orthotropic (Rauh’s ing an extended apex Agathis
model), sometimes profusely reiterating or sprout- 1b. Bark rough, exfoliating in horizontal strips or
ing from dormant buds in roots near the base of forming many irregular pustules; leaves ses-
stems. Foliage branchlets with or without terminal sile; seed cones ovoid-globose to globose; cone
buds. Leaves spirally arranged or sub­opposite, scale- scales with an extended, free apex 2
like and adnate or laminar and sessile or short petio- 2a. Bark forming many irregular pustules in ma-
late, imbricately covering the shoot or free and more ture trees; leaves opposite or subopposite,
or less distichously spreading, sometimes forming mostly linear, distichous or tetrastichous (in 2
4 ranks (Wollemia), more or less coriaceous, with or 4 ranks); pollen and seed cones terminal on
numerous parallel veins originating from basal di- first-order branches Wollemia
chotomies and few to numerous resin canals. Pollen 2b. Bark exfoliating in horizontal strips; leaves
cones axillary to leaves, solitary or in small clusters, spirally arranged, scale-like or triangular to
small or large, much elongating after anthesis and lanceolate, mostly equally divided around the
becoming cylindrical; microsporophylls numer- shoot; pollen and seed cones lateral on higher
ous, helically inserted, crowded, with imbricate or order foliage branches Araucaria
tesselate heads, each with 4–20 oblong pollen sacs
containing non-saccate pollen. Seed cones termi-
nal on long shoots (Wollemia) or lateral on short, Cephalotaxaceae Neger, Nadelhölzer: 23, 30. 1907
pedunculate, leafy shoots, solitary, erect, ovoid or (nom. cons.). Type: Cephalotaxus Siebold & Zucc.
subglobose, sometimes massive (Araucaria bidwillii ex Endl.
has the heaviest cones of all conifers), usually dis-
intergrating leaving the rachis on the tree. Bracts
Description
helically inserted on the rachis, much developed,
flattened, with a thickened distal margin and with See the genus description.
or without a terminal elongated cusp, forming the
bulk of the cone. Seed scales much reduced, axillary One genus: Cephalotaxus, with 8 species.
to and almost entirely fused with the bract, with or
without a small, free apical ligule, more or less en-
Distribution
closing a single, inverted seed, concrescent with the
seed scale or free, wingless or with a single wing or 2 As for the genus.
unequal wings. Seedlings with 2, sometimes deeply
divided, cotyledons. Number of chromosomes (di­
ploid) 2n = 26.
 Cupressaceae Gray, Nat. Arr. Brit. Pl. 2: 222, 225.
Taxonomic notes
1821. (nom. cons.), [“Cupressideae”]. Type: Cupressus
The family Cephalotaxaceae has been variously cir- L.
cumscribed by different authors, or included in the
Taxaceae s. l. This situation led Page (1990) to remark
Description
that “obscurity is probably the only aspect of the ge-
neric and family affinities … about which we can be Aromatic, resinous, evergreen or sometimes de-
totally sure.” He included Amentotaxus and Cephalo­ ciduous, monoecious or dioecious shrubs or trees,
taxus in Cephalotaxaceae, but his circumscription has ranging from diminutive prostrate shrubs to trees
not been generally accepted. Florin (1948) excluded exceeding 100 m. Bark fibrous or brittle, exfoliating
Taxaceae s. str. from ‘true’ conifers, but included in longitudinal strips or small plates. Branches erect 45
­Cephalotaxus, which he considered to “occupy an or spreading; foliage branches erect to pendulous,
isolated position among the recent representatives of initially covered with (decurrent) leaf bases until
this class.” Cephalotaxus appeared to Florin to have secondary growth replaces these with bark, terete,
several ovules aggregated into a cone, while the oth- (quadr)angular or more or less flattened, orthotropic
er genera lacked evidence of a cone, or the reduction or in more or less plagiotropic and often frond-like
of one. A number of morphological studies of both sprays. Buds mostly absent; (seasonal) abscission of
male and female reproductive organs have brought (pen)ultimate branchlets, not of single leaves. Foli-
the two groups more closely together, often inferring age leaves simple, usually polymorphic dependent
a more primitive state in Cephalo­taxus compared on life phase, with juvenile, transitional and mature
with Taxaceae s. str. The male cones in Taxaceae were forms, and on growth of foliage branches. Leaves on
interpreted by Wilde (1975) as reduced structures (pen)ultimate branchlets linear, needle- or scale-
derived from more clearly compound systems in like, spirally arranged, ternate or decussate (rarely
Cephalotaxaceae. These differences were confirmed quadrate) in mature plants, also in whorls of 4 in
in an elaborate study by Mundry (2000). The struc- juvenile plants and in some mature plants; decus-
ture and ontogeny of the female reproductive organs sate scale leaves pairwise dimorphic on flattened
however, are essentially similar in Cephalotaxus and branchlets. Leaf resin duct or cavity mostly single,
Taxaceae. The ‘cone’ Florin observed in Cephalo­ often with a dorsal gland in scale leaves. Leaf vascu-
taxus results from a swelling of the bracts subtending lar bundle single. Pollen cones small, terminal, rare-
the ovules, of which more in a single reproductive ly axillary, solitary or sometimes clustered in groups
unit are retained, at least to the pollination stage, in of 2–7, sessile on foliage branchlets or on dwarf
this genus than in the others. Other similarities are shoots, simple, deciduous. Microsporophylls spiral-
evident, but there are also some marked differences; ly arranged, ternate or decussate in congruence with
Chen & Wang (1990) found the numbers of free leaf phyllo­taxis on the cone-bearing shoot, small
nuclei in the female gametophyte to be extremely and thin, (sub)peltate. Pollen sacs (microsporangia)
high but variable (1024–4096) in ­Cephalotaxus, and abaxial, free, 2–10(–14) per microsporophyll, in one
relatively low and constant (256) in 4 of the 5 genera or two rows, longitudinally dehiscent, containing
of Taxaceae (Torreya was not investigated). Phylog- spheroidal, non-saccate pollen. Seed cones terminal,
enies reconstructed from molecular data (DNA nu- solitary or sometimes secondarily clustered, sessile
cleotide sequences) tend to place Cephalotaxus basal or pedunculate, simple or semi-compound, globose,
(as sister to) a clade with Taxaceae s. str. (e.g. Cheng ovoid or conical, deciduous or persistent. Bract-
et al., 2000; Quinn et al., 2002), but in the latter study scale complexes spirally arranged, in whorls of 3–4
the consensus tree shows a polytomy within a sin- or decussate in congruence with leaf phyllotaxis on
gle clade. This can be interpreted as evidence that the cone-bearing shoot, consisting of transformed
Cephalotaxus is best placed within Taxaceae (Quinn bracts enlarged with subsidiary intercalary growth,
et al., 2002), but the ‘total’ evidence is still ambigu- rarely with elements of ovuliferous scales, forming
ous. A solution of sorts could be to recognize this (semi-)woody or soft, imbricate or valvate, free or
genus as a subfamily Cephalotaxoideae, but until its (partially) fused, (sub)peltate, ovate or oblong cone
relationships are unambiguously resolved, it seems scales with or without an abaxially protruding bract
advisable to retain a monogeneric family Cephalo- tip. Ovules at base of bracts, axillary or sometimes
taxaceae. terminal, erect (or secondarily inverted), single to
 numerous. Seeds winged or unwinged, wings 1–3 ontogeny of cones using scanning electron micro-
per seed, often unequal in size and shape, derived scopes (SEM), there is no clear cut distinction, but a
from thin or thick and hard seed coat, seed often grade through both families. If there is a line of de-
with resin pits. Seedlings with 2–6(–9) cotyledons; marcation between the two positions, it lies within
germination epigeal. Chromosome number (dip- the traditional Taxodiaceae (see Farjon, 2005a for an
loid) = 22, sometimes tetra- or hexaploid (Sequoia overview). It was Eckenwalder (1976) who first dem-
sempervirens). onstrated in a comprehensive phenetic analysis of
morphological and other characters, that apart from
30 genera: Actinostrobus (3 spp.), Athrotaxis (3 spp.), Sciadopitys (formerly classified in Taxodia­ceae), no
Austrocedrus (1 sp.), Callitris (15 spp.), Calocedrus taxon assigned to Taxodiaceae was separable from
46 (4 spp.), Chamaecyparis (5 spp.), Cryptomeria (1 sp.), a larger group including taxa classified in the Cu-
Cunninghamia (2 spp.), Cupressus (15 spp.), Diselma pressaceae. Phylogenetic analyses, both of morpho-
(1 sp.), Fitzroya (1 sp.), Fokienia (1 sp.), Glypto­strobus logical data and of molecular (DNA) data, and of
(1 sp.), Juniperus (53 spp.), Libocedrus (5 spp.), combined data, all demonstrate that the genera of
Metasequoia (1 sp.), Microbiota (1 sp.), Neocallitrop- traditional Taxodiaceae constitute not a coherent
sis (1 sp.), Papuacedrus (1 sp.), Pilgerodendron (1 sp.), clade (group), but a ‘grade’ of taxa with on the whole
Platycladus (1 sp.), Sequoia (1 sp.), Sequoiadendron less advanced character states. These genera are also
(1 sp.), Taiwania (1 sp.), Taxodium (2 spp.), Tetra- predominantly monospecific, with in several cases
clinis (1 sp.), Thuja (5 spp.), Thujopsis (1 sp.), Wid- more species known from the fossil record. They
dringtonia (4 spp.) and Xanthocyparis (2 spp.); total therefore constitute a loose assembly of relict species
135 species. with more or less basal positions in the phylogeny of
the Cupressaceae. It is interesting to note that palae-
obotanists, who have been more hesitant than other
Distribution
systematists to abandon the concept of Taxodiaceae,
Cosmopolitan. Northern hemisphere: Macaronesia; are now also beginning to interpret Cupressaceae in
N Africa (including one locality in Central Sahara). the here accepted circumscription (e.g. Stockey et
Eurasia: Europe (including Iceland), SW Asia (excl. al. in Farjon, 2005a; Anderson et al., 2007). This is
driest deserts), Central Asia, Mongolia, Siberia, Rus- undoubtedly due to the fact that more fossil remains
sian Far East (including Kamchatka), Hindu Kush, are coming to light that appear to fill in the gaps cre-
Himalaya, China, Taiwan, Japan, Korea, Laos, Viet- ated by the extinctions.
nam. Australasia: New Guinea, Maluku (Moluccas),
Australia (including Tasmania). SW Pacific: New
Key to the genera
Caledonia, New Zealand. North America: Cana-
da, S Greenland, USA (including Alaska), Mexico. 1a. Leaves linear-lanceolate; margins serrulate;
Central America S to Honduras. Caribbean Islands. seed cones > 15 mm wide, composed of heli-
South America: S Chile and S. Argentina (Andes) to cally arranged, coriaceous scales each with 2–3
Tierra del Fuego. seeds Cunninghamia
1b. Leaves of a different shape, entire if linear; seed
cones if with helically arranged coriaceous
Taxonomic notes
scales, then <15 mm wide and 2 seeds per scale
The family Cupressaceae here includes the former 2
Taxodiaceae. The initial full description of Taxo- 2a. Leaves on lateral, short branchlets all linear, flat
diaceae by Warming (1884) included spiral phyllo­ and entire 3
taxis as a distinctive character; at that time 2b. Leaves on lateral, short branchlets at least
Metasequoia (with opposite phyllotaxis but in other in part scale-like or subulate, or a mixture of
characters similar to e.g. Sequoia and Taxodium in scale-like and linear leaves present on the same
Taxodiaceae) was unknown. Another distinction mature plant 5
claimed in earlier works was the position of ovules 3a. Leaves on lateral branchlets and cone scales
in developing cones: axillary to bracts (Cupressa­ decussate, lateral branchlets deciduous
ceae) or basal upon them (Taxodiaceae). In reality, Metasequoia
as has now been shown in detailed observations of
3b. Leaves on lateral branchlets and cone scales 11a. Leaves on lateral branchlets decussate, dimor-
helically arranged 4 phic, with differently shaped facials and later-
4a. Leaves scale-like on leading branches, linear als 12
and flat on lateral branchlets, lateral branchlets 11b. Leaves on lateral branchlets decussate or
evergreen; all cones near or at the end of foliage whorled (3–4), monomorphic or at least not
branchlets Sequoia clearly divided in facials and laterals 23
4b. Leaves (almost) all linear, flat or short acicu- 12a. Seed cones (sub)globose when closed, with
lar to subulate in one variety; lateral branch- cone scales of more or less equal length 13
lets (semi-)deciduous; pollen cones numerous 12b. Seed cones oblong when closed, with some
on leafless branchlets; seed cones on thicker paired cone scales longer than other scales 18
branches, often aggregate Taxodium 13a. Seed cones with 4(–5–6) subpeltate scales in 47
5a. Leaves on lateral foliage branchlets and cone decussate pairs 14
scales all helically arranged 6 13b. Seed cones with 8 or more peltate or subpeltate
5b. Leaves on lateral foliage branchlets and cone scales in decussate pairs 16
scales decussate, opposite or whorled 10 14a. Leaves weakly dimorphic, appressed; columella
6a. Leaves on lateral foliage branchlets scale-like as in seed cone absent 15
well as linear, lateral branchlets deciduous 14b. Leaves strongly dimorphic, lateral leaves
Glyptostrobus with free apex, or monomorphic linear leaves
6b. Leaves on lateral foliage branchlets scale-like or present; columella present Xanthocyparis
subulate or lanceolate, lateral branchlets ever- 15a. Small tree or shrub; leaves long decurrent, ad-
green 7 nate to branchlets except for the small apex;
7a. Leaves on lateral branchlets mostly scale-like; seed cones woody, > 10 mm, with 4–6 winged
seed cones ovoid, > 30 mm long with 30 or seeds Tetraclinis
more peltate scales; bark on trunk thick and 15b. Decumbent shrub; leaves short decurrent; seed
soft fibrous Sequoiadendron cones < 5 mm, with a single, wingless seed
7b. Leaves on lateral branchlets scale-like, ap- Microbiota
pressed, or subulate or lanceolate; seed cones 16a. Leaves on lateral branchlets lustrous green
< 30 mm long with less than 30 scales; bark above, with conspicuous, white stomatal bands
relatively thin and scaly 8 below; seed cones > 11 mm 17
8a. Leaves on lateral branchlets of cone-bearing 16b. Leaves green above, with ± obscure, greenish
trees all subulate, distinctly keeled; seed cones white stomatal bands below; seed cones < 12
squarrose; cone scales cuneate, with small teeth mm Chamaecyparis
above the recurved bract tip Cryptomeria 17a. Seed cones with 2(–3) pairs of larger scales and
8b. Leaves on lateral branchlets scale-like, subulate 2–3 pairs of reduced scales; leaves thick, facials
or lanceolate; seed cone scales without small and laterals equally long Thujopsis
teeth above the bract tip 9 17b. Seed cones with at least 4 pairs of larger, peltate
9a. Leaves subulate in younger trees only, eventu- scales, only the upper 1–2 pairs reduced; leaves
ally changing to scale leaves; seed cones ellip- thin, laterals exceeding facials Fokienia
soid to cylindrical, with thin cone scales; pollen 18a. Seed cones with 3 decussate pairs of scales, the
cones aggregate Taiwania proximal pair reduced, the middle pair spread-
9b. Leaves subulate or scale-like on different ma- ing and fertile, the long distal pair fused and
ture trees; seed cones (sub)globose, with ± sterile Calocedrus
thickened cone scales; pollen cones solitary 18b. Seed cones with 2 or 3–6 decussate pairs of
Athrotaxis scales, the middle and distal pairs spreading
10a. Seed cones berry-like, ± soft, indehiscent; cone and fertile, or the distal pair reduced and ster-
scales (almost) entirely fused; leaves scale-like ile 19
or acicular-linear Juniperus 19a. Seed cones with 2 decussate, fertile pairs of
10b. Seed cones not berry-like, (eventually) dehis- scales 20
cent; cone scales at least partly integer; leaves 19b. Seed cones with 3–6 decussate pairs of fertile
on late1al branchlets scale-like, rarely acicular and sterile scales 22
or linear 11
 20a. Seed cone scales with a large bract apex 29a. Valvate, thick cone scales subtended by ap-
Libocedrus pressed, thin sterile scales; leaf apices on ulti-
20b. Seed cone scales with a small bract apex 21 mate branchlets spreading and acute
21a. Facial leaves minute in comparison to laterals Actinostrobus
in all stages; bract apex nearly central on the 29b. Valvate, thick cone scales not subtended by
seed cone scale Papuacedrus sterile scales; leaf apices on ultimate branchlets
21b. Facial leaves slightly smaller than laterals; bract usually appressed or just free Callitris
apex subapical Austrocedrus
22a. Seed cone scales thin, the distal pair reduced
and connate; seeds winged Thuja Phyllocladaceae Bessey, Nebraska Univ. Stud. 7:
48 22b. Seed cone scales thickened, the distal pair well 325. 1907. Type: Phyllocladus Rich. ex Mirb.
developed and spreading; seeds unwinged
Platycladus
Description
23a. Leaves on lateral branchlets decussate 24
23b. Leaves on lateral branchlets whorled, usually See the genus description.
ternate 27
24a. Seed cones globose to ovoid, with > 6 peltoid One genus: Phyllocladus, with 4 species.
scales; seeds numerous, small, winged
Cupressus
Distribution
24b. Seed cones with a different shape and/or fewer
scales; seeds few to numerous, winged or nearly As for the genus.
wingless 25
25a. Shrub to very small tree (to 6 m); seed cones
Taxonomic notes
tiny, ca. 3 mm long, with 2 pairs of scales; maxi-
mal 2 seeds per cone Diselma The genus Phyllocladus was placed within the Po-
25b. Shrub to tall tree, seed cones > 10 mm long, docarpaceae as a subfamily Phyllocladoideae by
with 2 or more pairs of scales; > 2 seeds per Pilger (1903), but elevated to family rank by Bessey
cone 26 (1907), a publication apparently overlooked by Keng
26a. Seed cones subglobose when closed, with thick (1973), who proposed the family again. Since then,
woody scales and numerous seeds per cone; this family status has been controversial. Keng, in
leaves on older branchlets more or less spirally a series of papers re-published in Taipei (Taiwan)
arranged Widdringtonia as ‘A monograph of the genus Phyllocladus (Con-
26b. Seed cones ovoid-oblong when closed, with iferae)’ (Keng, 1963–79), emphasised the peculiar
thin woody scales and 3–4 seeds per cone; all phylloclades which function as true leaves and, er-
leaves decussate Pilgerodendron roneously, believed they were homologous with pro-
27a. Leaves all in alternate whorls of 4, seemingly in gymnospermous dichotomous branching systems,
8 rows, lanceolate; seed cones rare, with 8 only e.g. those of the Devonian genus Archaeopteris. We
2-3 mm wide scales Neocallitropsis now know that they are adaptive structures of much
27b. Leaves usually in alternate whorls of 3, scale- more recent origin, unique in conifers but evolved
like or sometimes acicular (juvenile form); seed independently in some angiosperms. Keng’s inter-
cones common, with 6, rarely with 8, > 3 mm pretation was criticised by Quinn (1987) and in a se-
wide scales 28 ries of subsequent papers (Chaw et al., 1997; Conran
28a. Mature type leaves long decurrent, adnate to et al., 2000; Quinn et al., 2002; Quinn & Price, 2003)
branchlets except for the (small) apex; seed researchers analysed various DNA sequence data to
cones with thick woody, valvate scales and a address the question of the phylogenetic relation-
± conical, hard columella 29 ship of Phyllocladus. The presented cladograms pre-
28b. Mature type leaves short decurrent, loosely dominantly show a clade ‘Podocarpaceae’ consisting
appressed or spreading; seed cones with thin of selected representative taxa, with Phyllocladus in
woody, widely spreading scales and a trigonal a basal position (sister to the other genera). The im-
to tripartite, soft columella Fitzroya plication inferred by Quinn and co-workers is that
Phyllocladus “is a true podocarp despite its highly
distinctive morphology.” If retained in the Podo­ resin canals and ray tracheids. Branching in rhyt-
carpaceae, its phylogenetic relationship as inferred mic pseudo-whorls on main stem and branches,
from these molecular data would obviously war- with apical dominance maintained or restored and
rant status as a subfamily. Whether one recognizes limited reiteration (Massart’s and Rauh’s models);
it instead as a family then depends on two criteria. branches terminating in seasonally dormant buds.
One is the acceptance or rejection of paraphyletic Leaves spirally inserted on long or short shoots and
taxa: when Phyllocladus and the Podocarpaceae solitary, or in fascicles of 1–8 surrounded by a sheath
are recognized in equal rank (both as family) and on dwarf shoots (Pinus), acicular-linear to long lin-
given the results of these analyses (phylogeny rep- ear, amphistomatic or hypostomatic, sometimes
resented by cladograms), the Podocarpaceae would epistomatic. Leaf anatomy in cross section with 1–2
be paraphyletic (i.e. a group not including all its vascular bundles encapsulated in a stele and 1-many 49
descendants) instead of monophyletic (including resin canals variously positioned in the mesophyll;
all its descendants). But the analyses remain lim- palisade parenchyma present or absent; hypodermis
ited; not all possible relatives were included. Based and epidermis usually well developed and differenti-
on the current information it is therefore appropri- ated. Pollen cones lateral on long shoots or apical on
ate to conclude that Phyllocladus is closely related short shoots, often grouped close together on long
to Podocarpaceae, but because of the limitations of shoots, axillary, solitary or clustered from a sin-
the molecular studies thusfar undertaken and the gle bud, catkin-like, deciduous. Microsporophylls
morphology, the Phyllocladaceae are for the present numerous, spirally arranged, (sub)peltate, with 2
retained as a separate family. The other criterion is abaxial, free pollen sacs (microsporangia) which
the evolution of morphological and biological char- are longitudinally dehiscent, containing bisaccate
acters. Apart from the phylloclades (which, being or monosaccate or non-saccate pollen. Seed cones
unique within conifers, provide no information woody, small to very large, erect or becoming pendu-
about phylogenetic relationships of members of lous, lateral on long shoots or apical on short shoots,
that group), there are some quite fundamental dif- often grouped together on long shoots, solitary in
ferences that would warrant recognition at a higher leaf axils, sometimes long persistent. Bracts spirally
taxonomic rank. These are seen in the reproductive arranged on a woody and integer rachis (breaking
morphology (Tomlinson et al., 1989), where Phyllo- up in Pseudolarix), remaining small or growing
cladus has an aril instead of an epimatium, and in with the cone; growing bracts apically differentiated
the different pollination mechanisms (Tomlinson et and often exserted. Seed scales axillary to and free
al., 1997). There are a suite of other characters which from bracts, both persistent or deciduous (Abies,
either “unite” Phyllocladus with Podocarpaceae ­Cedrus); seed scales flattened; apex rounded, more
(Quinn et al., 2002) or “separate” them (Page, 1990), or less acute (Pseudolarix), or differentiated (Pinus).
and each author tends to emphasize those in favour Ovules 2 per scale, inverted; seeds slightly flattened,
of their respective views on the matter. Taxon recog- usually winged (with 1 wing) but in some species
nition, despite claims to the contrary, is not a matter of Pinus effectively wingless due to rudimentary
to be decided in a purely formalized way (e.g. cladis- wings remaining attached to the seed scale. Seed-
tic analysis) and more evidence is needed to resolve lings with varying numbers of cotyledons (2–24, the
this question to the satisfaction of all. highest number in all plants). Germination epigeal
(hypo­geal in Keteleeria); young taproot of the seed-
ling with 1–2 resin canals in the vascular cylinder
Pinaceae Spreng. ex F. Rudolphi, Syst. Orb. Veg.: 35. (stele). Diploid number of chromosomes (2n) = 24
1830. Type: Pinus L. (in Pseudo­larix 44 and in Pseudotsuga menziesii 26).

11 genera: Abies (47 spp.), Cathaya (1 sp.), Cedrus

Description
(3 spp.), Keteleeria (3 spp.), Larix (11 spp.), Notho­
Monoecious evergreen or deciduous, resinous trees tsuga (1 sp.), Picea (38 spp.), Pinus (113 spp.), Pseudo­
or shrubs, sometimes to 100 m tall. Resin canals larix (1 sp.), Pseudotsuga (4 spp.) and Tsuga (9 spp.);
in wood, bark, leaves and seed cones. Wood with total 231 species.
adaxial parenchyma, with normal (non-traumatic)
 ous; seed cones erect from mostly pendulous
Distribution
branches Larix
Northern hemisphere (with one equatorial cross- 4b. Shoots monomorphic or weakly dimorphic;
over in N Sumatera). Eurasia & N Africa: Canary leaves spirally arranged, remote or dense, ever-
Islands, Mediterranean coast of N Africa, Europe green; seed cones spreading or pendulous from
(excl. Iceland & Ireland), Turkey, Syria, Lebanon, mostly plagiotropic branches 5
Caucasus, Central Asia, Siberia, Russian Far East 5a. Shoots weakly dimorphic, with (alternate) long
(including Kamchatka), Korea, Japan, Hindu Kush, and short growth; leaves in short growth parts
Himalaya, NE India, China, Taiwan, Indochina, of shoots in dense tufts; bract scales of seed
N Sumatera, Philippines. North America: Canada, cones small, included and simple acuminate
50 USA (including Alaska, excl. some midwestern Cathaya
states), Mexico. Central America S to Nicaragua; 5b. Shoots monomorphic; leaves equally distant on
Caribbean Islands (excl. Lesser Antilles). all (segments of) shoots; bract scales of cones
large, exserted and trilobate at apex
Pseudotsuga
Key to the genera
6a. Mature seed cones pendulous Tsuga
1a. Adult green leaves bundled with 2–5(–8) togeth- 6b. Mature seed cones erect 7
er on dwarf shoots (1 species with a single leaf 7a. Pollen cones in (umbellate) clusters from a sin-
on a dwarf shoot); seed cones biennial (rarely gle bud; seed cones on mostly long, leaved pe-
triennial), with distinction between each year’s duncles; cone rachis (slowly) disintegrating 8
growth apparent as an umbo and apophysis on 7b. Pollen cones solitary; seed cones on short, bare
each scale; seed held to the wing in a pair of peduncles or sessile; cone rachis persistent 10
claws Pinus 8a. Shoots strongly dimorphic; leaves spirally and
1b. Adult green leaves either solitary or in pseu- remotely arranged on long shoots, in dense
do-whorls of more than 10 together on short pseudo-whorls on short shoots, deciduous;
shoots; seed cones mostly annual (if biennial seed scales deciduous by rapid disintegration of
without umbo); seed held to the wing in a cup 2 the cone rachis Pseudolarix
2a. Seeds without resin vescicles; seed scales with a 8b. Shoots monomorphic or weakly dimorphic;
broad basis, persistent 3 leaves spirally arranged on long shoots, ever-
2b. Seeds with resin vescicles; seed scales with a green; seed scales (longer) persistent 9
narrow, petiolate basis, persistent or deciduous 9a. Seed cones large, (usually) more than 6 cm long
6 and 3 cm wide; leaves with narrowed, twisted
3a. Seeds easily separated from relatively large, base, 2–4.5 mm wide, on weakly developed pul-
narrow seed wing; pulvini on shoots very vini, leaving circular leaf scars after falling
strongly developed; leaves mostly amphisto- Keteleeria
matic or sometimes epistomatic; seed cones 9b. Seed cones small, (usually) 2.5–5 × 1.5–2.5 cm;
with rudimentary (or small) bract scales and leaves with oblique, petiolate base, 1–2 mm
with rudimentary scales on the pedunculate wide, on more developed pulvini, leaving no
base, pendulous at maturity Picea distinct leaf scars after falling Nothotsuga
3b. Seeds not separable from relatively small, broad 10a. Shoots strongly dimorphic; leaves spirally and
seed wing; pulvini on shoots weakly developed remotely arranged on long shoots, in dense
or absent; leaves epi/hypostomatic or epi-am- pseudo-whorls on short shoots; seed cones ma-
phistomatic with distinct primary stomatal turing in two years, disintegrating on tree with
bands; seed cones with small to very large bract deciduous seed scales Cedrus
scales and distinct, leaved peduncles, erect, 10b. Shoots monomorphic; leaves spirally arranged
spreading or pendulous at maturity 4 on long shoots; seed cones maturing in one sea-
4a. Shoots strongly dimorphic; leaves spirally son, with deciduous bracts and seed scales
and remotely arranged on long shoots, in Abies
dense pseudo-whorls on short shoots, decidu-
Podocarpaceae Endl., Syn. Conif.: 203. 1847. (nom. a hard, sclerified seed coat and wingless. Seedlings
cons.). Type: Podocarpus L’Hér. ex Pers. with 2 cotyledons, germination epigeal. Chromo-
some numbers (diploid) 2n = 18 to 38 (but not 28
and 32).
Description
Dioecious or sometimes monoecious, evergreen, 18 genera: Acmopyle (2 spp.), Afrocarpus (5 spp.),
slightly resinous trees or (dwarf) shrubs (one par- Dacrycarpus (9 spp.), Dacrydium (22 spp.), Falcati­
asitic on another member of the family). Bark of folium (6 spp.), Halocarpus (3 spp.), Lagarostrobos
trees thin, usually becoming scaly and exfoliating in (1 sp.), Lepidothamnus (3 spp.), Manoao (1 sp.),
flakes or strips. Tree trunks monopodial, commonly Micro­cachrys (1 sp.), Nageia (5 spp.), Parasitaxus
with orthotropic branching (Rauh’s model). Foliage (1 sp.), Pherosphaera (2 spp.), Podocarpus (97 spp.), 51
branches terminating in buds with small, leaf-like Prumnopitys (9 spp.), Retrophyllum (5 spp.), Saxe-
or more specialized scales (Podocarpus), or termi- gothaea (1 sp.) and Sundacarpus (1 sp.); total 174 spe-
nating in reduced leaves. Leaves helically arranged, cies.
rarely decussate (Microcachrys), appressed and
imbricate to spreading and remote, shapes highly
Distribution
variable, ranging from small, appressed scale leaves,
via thin acicular leaves, to dorsiventrally flattened, Pantropical, extending in the southern hemisphere
linear-lanceolate to broadly lanceolate, large leaves to cool temperate and in the northern hemisphere
(up to 34 × 9.5 cm); venation a single median strand to warm temperate latitudes. Africa: Sub-Saharan
or multiple parallel veins; texture soft and flexible to Africa only, in W Africa from Sâo Thomé, S Nigeria
coriaceous and more or less stiff; leaves amphisto- and Cameroon to Angola; in E and S Africa from
matic or hypostomatic. Pollen cones mostly simple, Ethiopia along the Afromontane system to the Cape;
axillary or terminal, solitary or clustered, in some Madagascar. Asia: S India (disjunct in Kerala) and
genera (Nageia, Prumnopitys) forming compound, from the E Himalaya and Assam through S China
racemose units (spikes), sessile or pedunculate, very to Japan; Taiwan, Indochina, Andaman and Nicobar
small to long cylindrical; microsporophylls spirally Islands, and Malesia. Australasia and SW Pacific:
arranged, with 2 abaxial, longitudinally dehiscent Australia (excl. the dry interior and NW), Solomon
pollen sacs; pollen bisaccate or trisaccate (rarely Islands, New Caledonia, Vanuatu, Fiji, Tonga, and
4–6), or non-saccate (Saxegothaea). Seed cones ax- New Zealand. Americas: From Mexico to Panama,
illary or terminally, solitary on naked or scaly pe- Caribbean Islands (excl. Bahamas & Turks Caicos
duncles, or sessile, composed of 2-many alternate Islands); South America along the Andean Cordill-
or helically arranged bracts; fertile bracts 1-many. era from Venezuela to S Chili, Venezuelan High-
Sterile and fertile bracts either fusing and forming a lands, SE and SW Brazil.
fleshy or succulent, often brightly coloured recepta-
cle, or remaining small as part of a small, compound
Key to the genera
cone and either becoming succulent and coloured
(Microcachrys) or leathery and dry (Pherosphaera, 1a. Leaves monomorphic, forming a distinct lami-
Saxegothaea). Seed scales single in the axil of a na (blade) wider than thick 2
bract, bearing a single initially erect but at pollina- 1b. Leaves monomorphic or dimorphic, at least in
tion more or less inverted ovule, in most genera (not part scale-like or more or less acicular, these
in Pherosphaera) much expanding after fertiliza- not or only slightly wider than thick 8
tion to enclose the seed partly or entirely, forming 2a. Leaves with multiple parallel veins Nageia
the epimatium. Epimatium leathery or fleshy, thin 2b. Leaves with a single (sometimes inconspicu-
or very thick, becoming succulent in some genera ous) midvein 3
that do not develop (or only imperfectly develop) a 3a. Seed cones forming an inflated, coloured, suc-
succulent receptacle, remaining green or variously culent receptacle below the seed(s) when ripe
coloured. Seeds (except Microcachrys, Pherosphaera Podocarpus
and Saxegothaea) single or sometimes 2, protrud- 3b. Seed cones much reduced, not or rarely slightly
ing well outside the transformed or reduced cone inflated, or consisting of numerous small, fer-
proper. Seeds proper ovoid, slightly flattened, with tile scales 4
 4a. Seed cones consisting of numerous small, fer- 15b. Epimatium surrounding the seeds completely,
tile scales Saxegothaea becoming swollen and succulent 17
4b. Seed cones much reduced, not or rarely slightly 16a. Leaves with well developed lamina always
inflated 5 present and dominant on all foliage branchlets
5a. Pollen cones aggregated in spikes Prumnopitys Acmopyle
5b. Pollen cones in clusters, sessile or very short 16b. Laminar leaves present or absent, not domi-
pedunculate, or solitary 6 nant (i.e. scale leaves predominate on some or
6a. Leaves in apparent opposite and decussate pairs most foliage branchlets) Dacrycarpus
due to twisted petioles turning the leaves in op- 17a. Leaves with well developed lamina always
posite directions Retrophyllum present and dominant on all foliage branchlets,
52 6b. Leaves alternate, not twisted in opposite direc- mostly falcate; seed cones forming a swollen re-
tions 7 ceptacle Falcatifolium
7a. Leaves with stomata on both surfaces (amphis- 17b. Leaves with well developed lamina absent or
tomatic) Afrocarpus present, but not dominant (i.e. scale leaves pre-
7b. Leaves with stomata only on the abaxial (lower) dominate on some or most foliage branchlets);
surface (hypostomatic) Sundacarpus seed cones much reduced Halocarpus
8a. Leaves monomorphic (only one type present)
9
8b. Leaves dimorphic (two distinct types present) Sciadopityaceae Luerss., Grundz. Bot.: 265. 1877,
14 [“Sciadopityeae”]. Type: Sciadopitys Siebold & Zucc.
9a. Small shrubs or dwarf trees with reddish scale
leaves and purple (pen)ultimate branchlets;
Description
seeds terminal, glaucous white Parasitaxus
9b. Shrubs or trees with green scale leaves on foli- See the species description.
age branchlets; seeds lateral or (sub)terminal,
usually not glaucous white 10 One genus: Sciadopitys, with 1 species.
10a. Dwarf shrubs, usually creeping, with decussate
scale leaves Microcachrys
Distribution
10b. Shrubs or trees with spirally arranged scale
leaves 11 As for the species.
11a. Seed cones very small, consisting of multiple
fertile (and some infertile) scales and ovules
(seeds) 12 Taxaceae Gray, Nat. Arr. Brit. Pl. 2: 222, 226. 1822.
11b. Seed cones much reduced, usually with only (nom. cons.). Type: Taxus L.
one fertile scale producing a single seed 13
12a. Always shrubs, trailing or upright
Description
Pherosphaera
12b. Trees (sometimes appearing shrubby with mul- Dioecious or rarely monoecious evergreen shrubs
tiple stems, especially when in cultivation) or trees, slightly resinous with resin in leaves (with
Lagarostrobos or without resin canals) and arils. Bark thin, exfo-
13a. Epimatium forming a yellowish green collar liating in strips. Branches spreading or ascending,
around the erect seed; always trees Manoao often with much reiteration; foliage branches termi-
13b. Epimatium basal, hidden from view by invert- nating in buds with small, imbricate scales. Leaves
ed ripe seed; (dwarf) shrubs or sometimes trees helically or decussately inserted, usually pectinately
Lepidothamnus arranged on plagiotropic branchlets, linear to lan-
14a. Leaves acicular, rarely scale-like, never with a ceolate, bifacially (dorsiventrally) flattened, with
distinct lamina Dacrydium a single median vascular bundle (midrib), coria-
14b. Leaves scale-like and/or with a distinct lamina, ceous, hypostomatic, with two separate bands of
never acicular 15 stomata. Pollen cones lateral on foliage shoots and
15a. Epimatium forming a collar around the base of axillary to foliage leaves, or aggregated in umbel-
the seeds 16 late clusters of (1–)3–6 racemes at or just below the
apex of vegetative shoots (Amentotaxus), subtended E USA to Florida; Mexico, Guatemala, Honduras, El
by small or large perular scales, short pedunculate, Salvador.
(sub)globose to short cylindrical. Microsporophylls
helically or decussately arranged (with secondary
Key to the genera
displacement forming 8 ranks), few in number (but
up to 60 in Torreya), radially symmetric (peltate) 1a. Leaves opposite-decussate; pollen cones in um-
or dorsiventrally compressed, with 2–8 pendulous, bellate clusters of (1–)3–6 racemes at or just be-
oblong pollen sacs either in radial placement or low the shoot apex Amentotaxus
abaxially, containing spherical, inaperturate pollen. 1b. Leaves alternate (spirally inserted); pollen
Seed bearing structures consisting of axillary dwarf cones solitary in the axils of leaves along the
shoots with decussate scales, branching 1–2 times, shoot (proximally, along the entire length or 53
usually strongly reduced, with terminal, erect ovules distally) 2
of which only a single one matures. Ovules at polli- 2a. Leaves usually not longer than 3.5 cm (rarely to
nation time around the base of the integument with 4.5 cm); aril surrounding the seed incompletely
a ring-like structure. Aril partially or entirely enclos- when mature, leaving the seed well visible 3
ing the seed, fleshy or succulent, becoming red, pur- 2b. Leaves usually longer than 4 cm and up to
ple, or yellow when ripe. Seeds oval to ovoid, slightly 12(–17) cm; aril surrounding the seed (almost)
flattened, with a hard, sclerified seed coat. Seedlings completely when mature, leaving the seed (vir-
with 2 cotyledons, germination epigeal. Number of tually) invisible 4
chromosomes (diploid) 2n = 22 (Amentotaxus, Tor- 3a. Foliage branchlets sub-whorled or subopposite;
reya) or 24. stomatal bands white; aril of seeds white when
ripe Pseudotaxus
Five genera: Amentotaxus (6 spp.), Austrotaxus (1 3b. Foliage branchlets irregularly alternate; sto-
sp.), Pseudotaxus (1 sp.), Taxus (10 spp.) and Torreya matal bands pale greyish green or pale yellow;
(6 spp.); total 24 species. aril of seeds red or sometimes orange to yellow
when ripe Taxus
4a. Midrib forming a narrow groove on the adaxial
Distribution
(upper) face of leaves; pollen cones aggregated
Western Eurasia & North Africa: Madeira; Atlas near the base of new foliage shoots; arils sur-
Mountains; Europe from Ireland and Portugal to rounding the seeds 12–15 × 7–9 mm, leaving the
S Scandinavia, European Russia and the Caucasus; seed apex visible Austrotaxus
Turkey, N Iran. Asia: Hindu Kush, Himalaya, Assam; 4b. Midrib on the adaxial face of leaves more or less
China, Korea, Russian Far East (Primorye), Japan, prominently raised; pollen cones forming dou-
Taiwan; Indochina; Malesia (Sumatera, Sulawesi, ble rows along the foliage shoots; arils 15–40 ×
Philippines). SW Pacific: New Caledonia. North & 10–25 mm, completely surrounding the seeds
Central America: British Columbia, Washington, Torreya
Idaho, W Montana, Oregon, California; E Canada,
TAXONOMIC TREATMENT OF GENERA AND SPECIES

Abies Mill., Gard. Dict., Abridg. Ed. 4, vol. 1. 1754. Type: Abies alba Mill. [Pinus
picea L.] (Pinaceae).
55
Abies is the classical Latin name for firs. Great Slave Lake to Newfoundland and Appalachian
Mountains in North Carolina. Eurasia and N Africa:
from Morocco through S and Central Europe to
Description
Turkey and Lebanon; NE Russia, Siberia and Central
Evergreen monoecious trees with a monopodial, Asia to Sakhalin and Japan; Sino-Himalayan moun-
straight, columnar trunk. Resin canals in bark, leaves, tain system and isolated spots in China, Taiwan.
and seed cones, normally not in wood unless trau-
matic. Branching in rhythmic pseudo-whorls on the
Taxonomic notes
trunk (Massart’s model), spreading horizontally, with
a dorsiventral symmetry (plagiotropic), branching The genus Abies is the second largest in Pinaceae
again with opposite shoots of lesser potential. (Sub-) (after Pinus). Liu (1971) recognized 39 species, while
terminal buds often extremely resinous. Leaves lin- Rushforth (1987) came to a total of about 55. In Far-
ear, more or less flattened, spirally arranged and usu- jon (1990) 46 species were recognized. The World
ally twisted at the petiolate base, set on small, circular Checklist & Bibliography of Conifers (Farjon, 1998,
depressions on the shoot which are clearly visible af- [2001]) recognized 49, resp. 48 species. Since the
ter detachment; with two conspicuous white stomat- monographic work of Liu, several new species have
al bands abaxially and sometimes scattered stomata been described, mainly from China; furthermore,
also on the adaxial side. Pollen cones usually below his treatment of several SW Chinese taxa failed to
seed cones in the tree, axillary, pendulous, solitary, recognize differences on the species level. On the
catkin-like, with spirally arranged, peltate micro- other hand, a number of species mentioned by
sporophylls; bearing two pollen sacs with bisaccate Rushforth have been given infraspecific status here,
pollen. Seed cones restricted to the upper part of the mainly at the subspecies level. Recently, a few new
crown, axillary, solitary, sessile or short pedunculate, species were described from Mexico, but it appears
erect, disintegrating at maturity when drying. Bracts that we have arrived at a phase where these reflect
and seed scales helically attached to a stout or slen- narrower circumscriptions rather than truly new
der rachis which remains on the branches. Bracts in discoveries.
mature cones exserted or sometimes hidden, with
a central cusp; cusp elongated or not. Seed scales
Synopsis
cuneate to reniform, with a pedicellate base. Seeds
held in a membranous cup, covering about 0.7 part The classification of the genus into sections and sub-
of the seed; membrane continued in a well devel- sections used here to key out the species in the ge-
oped, cuneate-triangular, persistent wing. Seedlings nus Abies is essentially that of Farjon & Rushforth
with (3–)4–8(–10) cotyledons. (1989), also used in my book Pinaceae (Farjon, 1990),
but with a few emendations. Most notable of these is
47 species. the removal of Abies kawakamii from Taiwan in sec-
tion Balsamea (subsection Laterales) and its tentative
placement in section Momi (subsection Homo­lepides)
Distribution
as a result of a molecular (nuclear DNA) analysis by
North America: (disjunct) from Yukon to Arizo- Xiang et al. (2004). This molecular analysis broad-
na and California into Mexico to Honduras; from ly supported the classification given by Farjon &
 ­ ushforth based on morphological data, but it has
R Species A. sachalinensis, A. fraseri,
demonstrated the anomaly of their placement of this A. koreana, A. nephrolepis, A. veitchii
species. The other changes made here involve a few Sect. Grandis Engelm. emend. Farjon & Rush-
additions and subtractions of species. The classifica- forth
tion given below is arranged according to hypotheti- Species A. grandis, A. concolor,
cal phylogenetic relationships, but in the keys to the A. ­durangensis, A. guatemalensis
species the sections and subsections are presented in Sect. Oiamel Franco
alphabetical order, with sections first and subsections Subsect. Religiosae (Matzenko) Farjon &
under these. Rushforth
Species A. religiosa, A. vejarii
56 Genus Abies Mill. Subsect. Hickelianae Farjon & Rushforth
Sect. Abies Species A. hickelii, A. hidalgensis
Species A. alba, A. cephalonica, Sect. Nobilis Engelm.
A. nordmanniana, A. nebrodensis, Species A. procera, A. magnifica
A. cilicica, A. ×borisii-regis
Sect. Piceaster Spach emend. Farjon & Rush-
Key to the sections and subsections
forth
Species A. pinsapo, A. numidica 1a. Bract cusps more than 2.5 cm long. Vegetative
Sect. Bracteata Engelm. emend. Sargent buds fusiform, 1–2 cm long, not resinous
Species A. bracteata Sect. Bracteata: A. bracteata
Sect. Momi Franco 1b. Bract cusps much shorter than 1.5 cm. Vegeta-
Subsect. Homolepides (Franco) Farjon & tive buds not fusiform, usually much less than
Rushforth, emend. 1 cm long, often resinous 2
Species A. homolepis, A. kawakamii, 2a. Seed cones very large (14–30 × 5–10 cm). Leaves
A. recurvata on vegetative shoots usually carinate, imbricate
Subsect. Firmae (Franco) Farjon & Rush- at base, lower leaves curving sideways, upper
forth leaves strongly assurgent Sect. Nobilis
Species A. firma, A. beshanzuensis 2b. Seed cones smaller, if longer than 15 cm then
Subsect. Holophyllae Farjon & Rushforth less than 6 cm wide. Leaves on vegetative shoots
Species A. holophylla, A. chensiensis, usually flattened 3
A. pindrow, A. ziyuanensis 3a. Seed cones narrowly cylindrical (ratio of length
Sect. Amabilis (Matzenko) Farjon & Rushforth to width greater than 2.5); rachis of cone coni-
Species A. amabilis, A. mariesii cal, slender 4
Sect. Pseudopicea Hickel emend. Farjon & 3b. Seed cones ovoid, conical or broad cylindrical;
Rushforth rachis of cone conical, cylindro-conical or fusi-
Subsect. Delavayianae Farjon & Rush- form, stout 8
forth 4a. Seed cones (10–)12–25(–30) cm long, ratio of
Species A. delavayi, A. fabri, A. for­ length to width usually 3 or more 5
restii, A. densa, A. spectabilis, A. far­ 4b. Seed cones 3–12 cm long, ratio length to width
gesii, A. fanjingshanensis, A. yuan­ usually less than 3 6
bao­shanensis 5a. Bracts exserted and reflexed (usually included
Subsect. Squamatae E. Murray in A. cilicica), with elongated cusp; cone apex
Species A. squamata obtuse or acutish (sometimes papillionate)
Sect. Balsamea Engelm. emend. Farjon & Sect. Abies
Rushforth 5b. Bracts always included, cusp short; cone apex
Subsect. Laterales Patschke emend. Farjon papillionate Sect. Piceaster
& Rushforth, emend. 6a. Seed cones small, (3–)4–8 × (1.5–)2–3(–4)
Species A. balsamea, A. lasiocarpa, cm, purplish (rarely greenish); bracts yellow-
A. sibirica brown. Leaves emarginate Sect. Balsamea 7
Subsect. Medianae Patschke emend. Far- 6b. Seed cones larger, (5–)7–12 × 3–5 cm, green, ol-
jon & Rushforth ive green or rarely bluish during the growing
 season. Leaves obtuse, acute or emarginate 2a. Leaves 1–2.2 cm long, 2–3.5 mm wide, rigid;
Sect. Grandis apex acute or obtuse, not emarginate. Seed
7a. Bracts exserted and reflexed; seed scales reni- cones small, 7–12 × 3–4 cm A. nebrodensis
form Subsect. Medianae 2b. Leaves (usually) longer than 2 cm, up to 2.5 mm
7b. Bracts included; seed scales cuneate-flabellate wide, acute to emarginate. Seed cones usually
Subsect. Laterales larger 3
8a. Seed cone rachis thick, fusiform or cylindro- 3a. Seed cones long and narrow (ratio length
conical; seed scales usually apically thickened. to width 3–4), width 3–5 cm. Leaves acute or
Shoots usually stout Sect. Pseudopicea 9 emarginate 4
8b. Seed cone rachis less thick, conical; seed scales 3b. Seed cones large, (10–)12–20 × 4–6 cm (ratio
thickest at or below the middle. Shoots usually length to width up to 3 at most). Leaves emar- 57
thin 10 ginate, on shaded, vegetative shoots pressed
9a. Bark exfoliating in large papery flakes forward, covering them A. nordmanniana
Subsect. Squamatae: A. squamata 4a. Leaves on shaded shoots emarginate, mostly in
9b. Bark different Subsect. Delavayianae pectinate arrangement with overlapping ranks.
10a. Bracts usually exserted with a broad cusp (but Young shoots pubescent 5
not in A. vejarii subsp. mexicana). Branchlets 4b. Leaves acute, rarely obtuse, more radially
purplish brown Sect. Oiamel 11 spreading. Young shoots glabrous
10b. Bracts included with a small cusp (which may A. cephalonica
be slightly exserted) or if exserted, cone ob- 5a. Leaves on the adaxial (upper) side lustrous
long-conical and green 12 green, without any stomata; apex of leaves on
11a. Leaves emarginate or obtuse; resin canals in the non-coning shoots slightly emarginate A. alba
leaves 4–10(–12) Subsect. Hickelianae 5b. Leaves on the adaxial side with a few stomata;
11b. Leaves acute; resin canals in the leaves 2 apex of all leaves entire A. ×borisii-regis
Subsect. Religiosae
12a. Young shoots uniformly pubescent; leaves
Key to the species of Section Amabilis
densely crowded above the shoot, directed for-
ward Sect. Amabilis 1a. Cones large, 9–15 × 5–8 cm. Leaves 2–3 cm long;
12b. Young shoots glabrous or weakly pubescent in buds ca. 5 × 4 mm A. amabilis
grooves; leaves not dense, more pectinate 1b. Cones smaller, 4–9 × 2–4.5 cm. Leaves short-
Sect. Momi 13 er (usually 1–2 cm long); buds small, 2–3 mm
13a. Seed cones oblong-conical; bracts strongly ex- long A. mariesii
serted Subsect. Firmae
13b. Seed cones ovoid-oblong to cylindrical; bracts
Key to the species of Section Balsamea,
included or cusps just exserted near the base of
Subsection Laterales
the cone 14
14a. Leaves 1.5–3(–3.5) cm long. Seed cones violet- 1a. Leaves on vegetative shoots mostly pectinate,
blue or purple, oblong-cylindrical to ovoid-­ a few above the shoot directed forward, short
oblong, 2.5–3.5(–4) cm wide and stiff. Seed cones (2.5–)5–8 × 2–3 cm
Subsect. Homolepides A. balsamea
14b. Leaves 2–5(–9) cm long. Seed cones yellowish 1b. Leaves on vegetative shoots mostly assurgent,
green to violet-blue, cylindrical or ovoid-cylin- short or longer, flexible. Seed cones mostly
drical, 3.5–6 cm wide Subsect. Holophyllae larger 2
2a. Leaves strongly assurgent, densely covering the
shoot; buds usually larger than 3 mm
Key to the species of Section Abies
A. lasiocarpa
1a. Bracts included or only slightly exserted. Shoots 2b. Leaves slightly assurgent, the lower leaves
mostly glabrous; leaves 2.5–4 cm long, with ob- spreading laterally, more remote, leaving the
tuse or slightly emarginate apex A. cilicica shoot visible; buds 2–3 mm long A. sibirica
1b. Bracts strongly exserted and reflexed 2
 Key to the species of Section Balsamea, Key to the species of Section Momi,
Subsection Medianae Subsection Firmae
1a. Seed cones with large, strongly exserted, re- 1a. Bracts oblanceolate, slightly exserted and not
flexed and light coloured bracts 2 reflexed. Buds large (max. 10 × 5 mm). Leaves
1b. Seed cones with smaller, less exserted and re- light green A. firma
flexed, usually darker coloured bracts 3 5b. Bracts spathulate, exserted and reflexed at the
2a. Seed cones very numerous, even on the lower upper margin. Buds smaller. Leaves dark green
branches, often crowded together, appearing A. beshanzuensis
soon on young trees; bracts leaving the apex of
58 the seed scales mostly free. Leaves on vegetative
Key to the species of Subsection Holophyllae
shoots emarginate A. koreana
2b. Seed cones absent from the lower branches, less 1a. Leaves on vegetative shoots entire 2
numerous; bracts very broad, covering most 1b. Leaves on vegetative shoots emarginate or
of the seed scales. Leaves on vegetative shoots bifid 3
mostly obtuse A. fraseri 2a. Seed cones oblong-cylindrical, 3–4.5 cm wide.
3a. Young shoots red-brown or brown, rarely grey- Leaves on leading and coning shoots strongly
brown. Leaves often strongly twisted at base; assurgent, acute or mucronate A. holophylla
cones usually ellipsoid-cylindrical. Bracts vari- 2b. Seed cones broader, 5–6(–7) cm. Leaves slightly
ously exserted A. sachalinensis assurgent and bifid, emarginate or sometimes
3b. Young shoots yellowish or greenish brown. obtuse A. pindrow
Leaves less twisted; cones cylindrical. Bracts 3a. Leaves 3–3.5 mm wide, 2–4.8 cm long, obtuse or
slightly exserted 4 rarely slightly emarginate. Ripe seed cones dark
4a. Young shoots densely yellowish pubescent. brown; bracts spathulate, with 9–10 mm wide
Leaves dark green above. Bracts slightly exsert- apex A. ziyuanensis
ed A. veitchii 3b. Leaves 2.5–3 mm wide, when wider, then long-
4b. Young shoots minutely pubescent in grooves. er than 4.5 cm, at least on vegetative shoots
Leaves light green above. Bracts entirely in- emarginate. Ripe seed cones (light) cinnamon-
cluded or only the cusps of the bracts exserted brown; bracts not spathulate A. chensiensis
A. nephrolepis
Key to the species of Subsection Homolepides
Key to the species of Section Grandis
1a. Leaves spreading pectinately or at least parted
1a. Leaves very thin, less than 2 mm wide, very above the shoot A. homolepis
flexible 2 1b. Leaves crowded and curved upwards or some-
1b. Leaves thicker, 2–3 mm wide, more rigid 3 times reflexed above the shoot 2
2a. Young shoots (light) greenish to reddish brown. 2a. Leaves on vegetative shoots 1.5–2 mm wide,
Leaves 1.2–2 mm wide, emarginate or obtuse. curved upwards A. kawakamii
Bracts slightly exserted A. guatemalensis 2b. Leaves on vegetative shoots 1.9–2.5 mm wide,
2b Young shoots dark purplish red or red-brown. recurved or reflexed A. recurvata
Leaves 1–1.6 mm wide, slightly acute or obtuse.
Bracts included A. durangensis
Key to the species of Section Nobilis
3a. Leaves green or glaucous green, stomata on
both sides, leaf apex entire A. concolor 1a. Bark of old trees (in their native habitat!) grey-
3b. Leaves dark glossy green above, stomata on one ish brown. Bracts of seed cones far exserted and
side only, leaf apex emarginate A. grandis strongly reflexed, covering much of the seed
scales. Leaves partly grooved above A. procera
1b. Bark of old trees (dark) red-brown. Bracts of
seed cones included or slightly exserted and re-
flexed. Leaves not grooved A. magnifica
 3b. Bracts gradually ending in a short cusp, not
Key to the species of Section Oiamel,
or slightly exserted. Young shoots yellowish
Subsection Hickelianae
brown A. densa
1a. Bracts of seed cones exserted A. hickelii 4a. Seed cones green or yellowish green in the
1b. Bracts of seed cones included A. hidalgensis growing season; bracts exserted and reflexed
A. yuanbaoshanensis
4b. Seed cones purple or purplish blue in the grow-
Key to the species of Section Oiamel,
ing season; bracts exserted or sometimes in-
Subsection Religiosae
cluded, not reflexed 5
1a. Seed cones (8–)10–16 × 4–6 cm; bracts strongly 5a. Young shoots usually purplish, reddish, or dark
reflexed, partly covering the seed scales. Leaves orange-brown 6 59
pectinately arranged, usually 1.5–3 cm long, 5b. Young shoots yellowish brown, mostly gla-
leaving the shoot visible from above brous A. fabri
A. religiosa 6a. Seed cones small, 5–9 × 3–4 cm; bracts with
1b. Seed cones 6–12(–15) × 4–6(–7?) cm; bracts not rounded or emarginate apex ending in a small,
reflexed. Leaves spreading but not pectinate, abrupt cusp 7
usually 1–2 cm long, covering the shoot above 6b. Seed cones 6–10(–14) × 4–5(–6) cm, or if less
A. vejarii than 8 × 4 cm, with differently shaped bracts
A. forrestii
7a. Leaves 1–2.5 cm long, rarely up to 4.3 cm, very
Key to the species of Section Piceaster
densely set in overlapping, pectinate rows.
1a. Leaves carinate, obtuse or slightly acute, Bracts mostly included or slightly exserted,
amphistomatic,very rigid and radial. Buds res- with very short cusps A. fanjingshanensis
inous. Seed cones 9–14 × 3–4 cm A. pinsapo 7b. Leaves (1–)1.5–3(–4.5) cm long, especially on
1b. Leaves flattened, weakly amphistomatic, on shaded shoots, less dense. Bracts exserted, with
shaded shoots less radial. Buds not resinous. much longer cusps A. fargesii
Seed cones usually larger, 12–18 × 4–6 cm
A. numidica
Abies alba Mill., Gard. Dict., ed. 8: Abies No. 1.
1768. Type: “Habitat in Alpinus Helvetiae, Sueviae,
Key to the species of Section Pseudopicea,
Bavariae, Scothiae”, Herb. Clifford 449 Abies 2
Subsection Delavayianae
(lectotype BM). Fig. 1
1a. Seed cones large, usually 10–17 × 4–7 cm; bracts
included. Leaves pectinately arranged, 2.5–6 Abies alba Mill. subsp. apennina Brullo, Scelsi &
cm long, 2.2–3.5 mm wide, bifid or emarginate, Spampinato, La Vegetazione dell’ Aspromonte: 41.
with slightly recurved margins A. spectabilis 2001.
1b. Seed cones smaller, if larger than 10 × 5 cm
usually with exserted bracts. Leaves spreading
Etymology
more radially, shorter or with strongly revolute
margins 2 The species epithet means ‘white’ and perhaps refers
2a. Leaves with (strongly) revolute margins; sto- to the light grey bark.
matal bands (partly) hidden, niveous white 3
2b. Leaves not revolute or with only very slightly
Vernacular names
recurved leaf margins, stomatal bands entirely
visible, white or greyish white 4 Silver fir; Sapin pectiné (French); Weisstanne (Ger-
3a. Bracts abruptly ending in an elongated, awl- man)
like cusp, usually exserted (at least with the
long cusp). Young shoots dark red-brown
Description
A. delavayi
Trees to 55(–60?) m tall, d.b.h. to 2–2.5 m; trunk mo-
nopodial, straight, columnar, terete; crown conical,
 but flat topped (“stork’s nest”) in most old trees. Bark
Ecology
in young trees greyish, smooth, in old trees greyish
brown, light or dark, becoming scaly and fissured Primarily a montane species, occurring between
below. Branches of first order horizontal, descend- 500 and 1500 m a.s.l., but as low as 300 m in the
ing, ascending towards the top of the tree; branches Bavarian Forest and up to 1950 m in the Pyrenees.
of second order slender, spreading horizontally. Soils are usually well drained sandy loams from
Branchlets slender, flexible, stout and firm on cone silicate rocks or limestone derivatives. The climate
bearing branches, yellowish grey or grey, nearly is cool temperate, comparatively humid (precipita-
smooth, striated, with short, yellowish or brown pu- tion often >1000 mm/year), with abundant snowfall
bescence or occasionally glabrous; leaf scars broadly but moderately low temperatures in the winter. The
60 elliptic or circular. Vegetative buds ovoid or conical, species often forms large forests, either pure, mixed
ca. 6 × 4 mm, not resinous or slightly resinous espe- with other conifers (Picea abies, locally Pinus sylves-
cially on cone bearing branches; bud scales broadly tris) or mixed with broad-leaved trees (Fagus sylvat-
ovate, obtuse, brown, persisting several years. Leaves ica), in a belt between deciduous forest in the valleys
spirally arranged, on vegetative branches pectinate and coniferous forest composed of other species of
in two lateral sets, on leading shoots radial, on con- Pinaceae towards the tree limit. Of the European na-
ing shoots radial and assurgent, (1.2–)1.5–3(–3.5) cm tive conifers, Abies alba is most capable of compet-
long, 1.5–2.6 mm wide, (slightly) twisted at base, ing with Beech (F. sylvatica) at altitudes where the
linear, flattened, grooved and lustrous dark green latter becomes less vigorous and is the first of the
on the adaxial (upper) face, whitish green below; conifers to appear among them (Ellenberg, 1988).
apex slightly emarginate (obtuse or acute on con-
ing shoots). Stomata in two bands separated by a
Conservation
midrib on the abaxial leaf face. Pollen cones lateral,
crowded, 2 cm long, greenish yellow with purple red IUCN: LC
microsporophylls. Seed cones lateral, erect; pedun-
cles short, slightly curved, cylindrical, with conical
Uses
or obtuse apex, 10–15(–20) cm long, 3–5(–6) cm
wide, yellowish green when immature, sometimes White fir is an important timber tree in western and
with purple tinge, ripening to light brown or red- central Europe, where most forests are semi-natural
dish brown; cone rachis persistent, narrowly coni- and managed with a view to encourage fir regen-
cal, dark brown. Seed scales cyathiform, the upper eration and growth at the expense of competitors.
ones more cuneate, length × width at mid-cone It is not very successful as a plantation forestry tree
2.5–3 × 2.5–3.5 cm; surface smooth, pulverulent es- outside its natural areas of occurrence; one cause of
pecially on the exposed part; upper margin entire; this may be damage done by insect pests, which may
base long pedicellate. Bracts linear-spathulate, with be more prolific in monocultures and in areas with
long, caudate cusp, 2.5–3.5(–4) cm long, exserted, mild winters such as in the British Isles. Its most fa-
slightly recurved at maturity. Seeds cuneate, angular, mous use in the past was for the masts of 17 th century
7–9 mm long, yellowish brown; seed wings cuneate, ships. Most of its wood today is used for plywood
10–15 mm long, reddish yellow. and veneer as it is evenly grained, light, and easily
worked. Minor uses are for soundboards in musical
instruments, boxes, wood carving, and sometimes
Distribution
for joinery. Distillation of the leaf essential oils has in
Europe: From the Pyrenees to the Carpathians, Italy the past been used to produce remedies for sprains
(Apennines) and in the Balkan Peninsula to Bul­ and bruises. In Britain it was used as a Christmas
garia and N Greece. tree in the 19th century when Prince Albert popular-
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SK ized the tradition; its use has declined since in favour
GER HUN POL SWI 12 COR FRA-FR SPA-AN SPA-SP of the more quickly and cheaply produced Norway
13 ALB BUL GRC ITA-IT ROM TUE YUG-BH YUG- spruce. In horticulture White fir is uncommon and
CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 mostly restricted to arboreta; a number of cultivars
UKR-UK are known but few are common in the trade.
Abies amabilis Douglas ex J. Forbes, Pinetum recurved when ripe, length × width at mid-cone
Woburn.: 125, t. 44. 1839. Picea amabilis Douglas 2–2.8 × 2.5–3.5 cm; surface smooth, often densely
ex Loudon, Arbor. Frut. Brit. 4: 2342. 1838. covered with resin, puberulous on both surfaces;
Type: A collection made by David Douglas, not upper margin entire, strongly incurved; base long
designated. Fig. 2 pedicellate. Bracts obovate-oblong, 1–1.5 cm long,
included, with very short cusps. Seeds oblong, 10–12
mm long, light brown; seed wings quadrangular,
Etymology
broad, 18 × 20 mm, yellow or pale brown.
The species epithet means ‘lovely’ and was coined by
its discoverer David Douglas.
Distribution
61
Pacific coast region of W North America, from ex-
Vernacular names
treme SE Alaska to N California.
Pacific silver fir, Lovely fir, Cascade fir TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL

Description Ecology
Trees to 45–80 m tall, d.b.h. to 2–3 m; trunk mo- Abies amabilis occurs from sea level near the coast
nopodial, straight, columnar, massive, terete; crown to 330 m a.s.l. in SE Alaska, in Oregon from 250 m
narrowly conical or irregular in old trees. Bark in to 1830 m a.s.l. on the western slopes of the Cascade
young trees smooth, ash grey or whitish, with resin Range. It grows on different mountain soils, usually
blisters, in old trees deeply furrowed, brown, divided of glacial origin and acidic. The climate is extremely
into small plates. Branches of first order horizontal, wet maritime, with 1500 to 4000 mm annual pre-
relatively short, thick, curved downward on lower cipitation, much of it as snow. It is a constituent of
part of the trunk; branches of second order short, the mixed coniferous forests with among other coni-
curved, spreading. Branchlets slender, light green- fer tree species Tsuga heterophylla, Picea sitchensis,
ish brown or greyish, changing to reddish brown, Pseudotsuga menziesii, Thuja plicata, Chamae­cyparis
ridged and grooved, remotely or more densely pu- nootkatensis, Abies grandis, A. magnifica; and with
bescent with light hairs; leaf scars circular, reddish. A. lasiocarpa and Tsuga mertensiana at higher eleva-
Vegetative buds subglobose, ca. 5 × 4 mm, very tions, but unlike the latter two not reaching the tree
resinous; bud scales dark purple, persisting sev- line.
eral years. Leaves spirally arranged, on vegetative
branches the lower ones pectinate, the upper ones
Conservation
directed forward, covering the branches, on coning
shoots radial, curved, the upper ones assurgent, 2–3 IUCN: LC
cm long, 1.5–2 mm wide, twisted at base, linear, flat-
tened, grooved and dark lustrous green above, two
Uses
silvery white lines below; apex slightly notched or
obtuse. Stomata in two bands separated by a mid- In the timber industry no distinction is made be-
rib on the abaxial (lower) leaf face, none or a few tween this species and Western hemlock as both
on the adaxial side. Pollen cones lateral in leaf axils conifers have similar wood properties. This wood
mostly solitary but numerous, 1–1.5 cm long, with is in use for various construction applications such
red microsporophylls. Seed cones lateral, erect, as plywood, veneer, sub-flooring and sheathing. It
mostly towards the ends of branches on short pe- contains little or no resin and is light in colour and
duncles, ovoid-cylindrical or oblong, with obtuse or easily worked. Together with Western hemlock,
papilliform apex, 9–15 cm long, 5–8 cm wide; purple substantial quantities of wood go to the kraft pulp
or rarely greenish when immature, purplish brown industry. As an ornamental tree it is uncommon,
when maturing, turning brown when ripe; cone performing only in cool and wet maritime climate
rachis persistent, cylindric conical, dark or light such as prevails in the west of Scotland.
brown. Seed scales cyathiform-flabellate or cuneate,
 Abies balsamea (L.) Mill., Gard. Dict., ed. 8: Abies long, entirely included or sometimes exserted and
No. 3. 1768. recurved. Seeds cuneate, 3–4 long, dark grey; seed
wings cuneate, oblique at apex, 7–10 mm long, grey-
ish brown.
Etymology
The species epithet refers to the ‘balsam’ i.e. its corti-
Distribution
cal resin which has medicinal properties.
Canada, North Central and NE USA: south to Vir-
ginia.
Vernacular names
TDWG codes: 71 ABT MAN SAS 72 LAB NBR NFL-
62 Balsam fir, Canadian fir; Sapin balsamier (French) NE NFL-SP 74 IOW MIN WIS 75 CNT MAI MAS MIC
NWH NWJ NWY PEN VER WVA 78 VRG
Description
Ecology
Trees to 25 m tall, d.b.h. to 1 m; trunk monopodial,
straight, columnar, terete; crown conical. Bark in Abies balsamea occurs from lowland plains to up-
young trees pale grey, smooth, with prominent resin land hills and mountains in the vast boreal forest
blisters, in old trees dark greyish brown or blackish, of North America, from sea level to 1200 m a.s.l. in
flaking. Branches of first order long and slender, in West Virginia. It is most common on usually pod-
whorls and horizontally spreading; branches of sec- zolized moderately acid soils in silt or sand. In some
ond order spreading horizontally. Branchlets slen- areas it may also grow on wet, peaty soil. The climate
der, pale yellowish green, becoming ash grey tinged is cold continental in the interior, cool maritime in
with red, smooth, shortly pubescent with dark grey the eastern part of the range, with precipitation be-
hairs; leaf scars circular to ellipsoid. Vegetative buds tween 250 and 1250 mm and very cold winters. The
ovoid globose, ca. 5 × 4 mm, resinous, with resin cov- growing season ranges from 80 days in the interior of
ering the scales entirely; bud scales dark green with Canada to 180 days in the Appalachian Mountains.
purple tinge, persisting several years. Leaves spirally It is a constituent of coniferous forests with Picea
arranged, mostly pectinate on sterile branches, the spp., Pinus strobus, Tsuga canadensis and sometimes
upper ones directed forward, assurgent on con- Pinus banksiana, or it grows mixed with broad-
ing shoots, 1.5–2.5 cm long, 2 mm wide, twisted at leaved trees such as Populus tremuloides, Betula spp.
base, linear, flattened, dark green above, two white and, further south, Acer spp., Fagus grandifolia and
bands separated by green midrib below; apex obtuse Betula alleghaniensis. Taxus canadensis is the most
or emarginate (acute or acuminate on cone bearing common conifer shrub in these mixed forests.
branches). Stomata on the adaxial (upper) surface
scattered along a medial groove, increasing towards
Uses
apex, on lower surface in two bands separated by a
midrib. Pollen cones lateral, in leaf axils, pendant, Balsam fir is an economically important conifer.
crowded and numerous, 0.6 cm long, yellow, with Its wood, although of modest size, is used in light-
purple microsporophylls. Seed cones lateral, erect; frame construction and for pulpwood. It is also pop-
peduncles very short; shape cylindrical, narrowing ular as a Christmas tree and is one of the top three
towards an obtuse apex, (2.5–)5–8 cm long, (1.5–)2–3 species grown for this purpose in E North America.
cm wide, green, tinged with purple, or violet-purple The fragrant needles are partly responsible for this
when immature, ripening to (purplish) brown; cone popularity, they are also used to stuff pillows sold as
rachis persistent, narrowly conical, blackish brown. souvenirs in New England. Canada balsam, the aro-
Seed scales cuneate, flabellate or rhombic-orbicular, matic and soft terpenoid resin collected from blisters
length × width at mid-cone 1.2–1.4 × 1.4–1.7 cm; in the bark, is especially important in Quebec. Its
surface smooth, striate on lower part, pubescent medicinal properties were known to Native Ameri-
with brown hairs on exposed part; upper margin cans, who used it as an antiseptic wound dress-
entire, strongly curved inward; base pedicellate. ing as well as internally for various ills. Lewis and
Bracts broad ligulate, with a short cusp, 0.6–1 cm Clark had it in their medicine box on their ­famous
­ verland expedition to the Pacific Ocean in 1804–
o
Taxonomic notes
05. In modern Western society its medicinal use has
been replaced by other salves; the resin is now used This fir has been interpreted by Liu (1971) as a hy-
to seal microscopic glass slides with biological pre- brid between A. balsamea and A. fraseri; this tax-
parates. In horticulture, Balsam fir is less valued; this onomy was accepted in my book Pinaceae (Farjon,
fir is apparently short lived when planted in gardens 1990) but is here rejected in favour of an infraspe-
and only a few dwarf cultivars are known. cific taxon under A. balsamea, with which it shares
more character states. Only the smaller cones with
2 varieties are recognized: exserting but small bracts remind of A. fraseri and
a hybrid origin of this variety has not been demon-
Abies balsamea (L.) Mill. var. balsamea. Pinus strated genetically. 63
balsamea L., Sp. Pl. 2: 1002. 1753. Type: “Habitat in
Virginia, Canada”, leg. ign. LINN 1135.14 (lectotype
Distribution
LINN).
USA: Virginia, West Virginia.
TDWG codes: 75 WVA 78 VRG
Description
Seed cones 5–8 cm long, 2–3 cm wide, green with
Conservation
a purplish tinge when young, brown at maturity;
bracts included or only the cusps exserting. IUCN: LC

Distribution
Abies beshanzuensis M. H. Wu, Acta Phytotax. Sin.
Canada, North Central and E USA: south to Vir- 14 (2): 16, t. 1, f. 1. 1976. Abies fabri (Mast.) Craib
ginia. var. beshanzuensis (M. H. Wu) Silba, Phytologia 68:
TDWG codes: 71 ABT MAN SAS 72 LAB NBR NFL- 13. 1990. Type: China: Zhejiang, Tung-Kung Mts.,
NE NFL-SP NSC ONT PEI QUE 74 IOW MIN WIS 75 Baishan-zu, M. H. Wu 7511 (holotype PE).
CNT MAI MAS MIC NWH NWJ NWY PEN VER WVA
78 VRG
Etymology
The epithet refers to the name of the mountain on
Conservation
which it was found.
IUCN: LC
Vernacular names
Abies balsamea (L.) Mill. var. phanerolepis Fernald,
Rhodora 11: 201. 1909. Abies intermedia Fulling, Baishan fir; Baishanzu lengshan (Chinese)
Castanea 1: 93. 1936; Abies balsamea (L.) Mill. f.
phanerolepis (Fernald) Rehd., Man. Cult. Trees, ed.
Description
2: 16. 1940; Abies ×phanerolepis (Fernald) T. S. Liu,
Monogr. Gen. Abies: 316. 1971; Abies balsamea (L.) Trees to 15 m tall, d.b.h. to 0.5 m; trunk monopo-
Mill. subsp. phanerolepis (Fernald) E. Murray, dial, straight, columnar, terete; crown broad, coni-
Kalmia 12: 18. 1982. Type: Canada, Quebec, Percé cal or flat topped. Bark of young trees whitish grey,
Mt., M. L. Fernald & J. F. Collins 860 (holotype GH). smooth, of old trees longitudinally fissured and scaly.
Branches of first order long, spreading horizontally,
ascending towards the top of the tree; branches of
Description
second order spreading and overlapping. Branchlets
Seed cones 2–5.5 cm long, 1.5–2 cm wide, violet-pur- stout, firm, light yellow, turning to greyish brown,
ple when young, greyish or blackish brown at matu- ridged and grooved between the leaves, glabrous or
rity, brown pubescent; bracts exserting, with long, sparsely pubescent; leaf scars circular or ovate ob-
reflexed cusps. long. Vegetative buds ovoid or conical, (slightly?)
resinous; bud scales triangular ovate, light yellow-
 ish brown, slightly keeled, persisting several years. leaved trees, e.g. Castanopsis spp., Fagus lucida,
Leaves spirally arranged, the lower ones pectinate, Quercus spp., Acer spp., Magnolia cylindrica and
the others directed obliquely forward, the short- Lithocarpus hancei. The angiosperm trees are domi-
est upper leaves recurved, parting above, on con- nating the present site of Abies beshanzuensis. Also
ing shoots crowded and assurgent, (1–)1.5–3.5(–4.2) reported is Cryptomeria fortunei; this is an invalid
cm long, 2.5–3.5 mm wide, twisted at base, ligulate name for C. japonica and not indigenous in China
linear or oblanceolate, flattened, shining dark green (Farjon, 1999). Its presence indicates secondary for-
above, two silvery white bands below; apex emar- est, as is also borne out by the photographs I have
ginate or bifid. Stomata absent on the adaxial (up- seen of the area.
per) surface, in two broad bands divided by a midrib
64 below. Pollen cones lateral, in leaf axils, 2–2.5 cm
Conservation
long, yellowish with red microsporophylls. Seed
cones lateral, erect; peduncles short and thick; shape Known from only a few mature individuals in the
oblong-conical, narrowing towards a truncate apex, wild, without natural regeneration, in degraded an-
(6–)7–11(–12) cm long, (3–)3.5–4(–4.5) cm wide, pale giosperm woodland. According to an account by
yellowish green when immature, turning light yel- Dudley (1988), in 1987 only three individual trees
lowish brown, ripening to light brown. Seed scales were left (after three other specimens had been re-
flabellate, auricled at base, length × width at mid- moved to Beijing Botanic Gardens and subsequently
cone 1.8–2.5 × 2.5–3 cm; surface striate, sparingly died there) of a population at discovery in 1963 of
pubescent (?); upper margin entire or slightly erose; only seven individuals, of which four were flower-
base pedicellate. Bracts spathulate, with reflexed up- ing and coning at that time. This species was taken
per margin, 2–2.5 cm long (cusps ca. 1 mm), exsert- into cultivation from cuttings at a forestry station
ed, recurved. Seeds oblong-obovate, length × width in Qingyuan, S Zhejiang, China, as a graft on Abies
6–9 × 3–4 mm; seed wings cuneate, oblique, length firma rootstock.
× width 6–10 × 5–9 mm, light brown. IUCN: CR (D)

Taxonomic notes Uses

This species resembles A. firma in several charac- No uses other than recent attempts to grow it in cul-
ters. Wu (1976) related this species to A. fabri on the tivation have been reported.
grounds of its recurved cone bracts, but both col-
our and shape of the cones are quite different and
resemble the cones of A. firma, as do the broad, flat Abies ×borisii-regis Mattf., Notizbl. Bot. Gart.
and emarginate leaves. Hence, Farjon & Rushforth Berlin-Dahlem 9: 235. 1925. Abies cilicica (Antoine
(1989) classified it in the section Momi, subsection & Kotschy) Carrière var. borisii-regis (Mattf.) Silba,
Firmae. Phytologia 68: 11. 1990; Abies alba Mill. subsp.
borisii-regis (Mattf.) Kozuharov & N. Andreev,
Opred. Vissh. Rast. Bulg.: 785. 1992. Type: Greece,
Distribution
Mt. Olympus, Monastery Hagios Dionysius, 1899,
China: Zhejiang (Mt. Baishan-zu NE of Qingyuan in P. E. E. Sintenis 1870 (lectotype, B).
the Tung-Kung Range at 27° 45´ N, 119° 11´ E).
TDWG codes: 36 CHS-ZJ Abies alba Mill. var. acutifolia Turrill, Kew Bull.
Misc. Inf. 1925: 34. 1925.
Abies borisii-regis Mattf. var. pungenti-pilosa Viguié
Ecology
& Gaussen, Bull. Soc. Hist. Nat. Toulouse 58: 279.
On a medium high mountain in the maritime SE of 1929.
China, with warm summers and cool, moist win-
ters (annual precipitation ca. 1250 mm), where it
Etymology
is found between 1500–1700 m a.s.l. It grows there
with other conifers, such as Tsuga chinensis, Cepha- This species was named after King Boris of Bulgaria
lotaxus chinensis and Taxus chinensis, and broad- (reigned 1918–1943).
 pus in Greece, and it is treated as such in several
Vernacular names
handbooks of conifers. Liu (1971) was of the opinion
King Boris fir, Bulgarian Fir, Macedonian fir that these trees can be best regarded as natural hy-
brids between A. alba and A. cephalonica. Although
often more similar to A. alba, most trees do share
Description
minor characters of both. More recent research into
Trees to 40 m tall, d.b.h. to 1.5 m; trunk monopo- hybridization (Mitsopoulos & Panetsos, 1987) has
dial, straight, columnar, terete; crown conical, but shown that introgression of A. alba genes into an-
flat topped (“stork’s nest”) in most old trees. Bark cient populations of A. cephalonica in the southern
of young trees greyish, smooth, in old trees greyish Balkans, due to southward migration of the former
brown, light or dark, becoming scaly and fissured during glacial stadia of the Pleistocene, is the likely 65
below. Branches of first order horizontal, descend- origin of A. ×borisii-regis. This process has led to
ing, ascending towards the top of the tree; branches more or less stable, true-breeding populations, but
of second order slender, spreading horizontally. has also reduced A. cephalonica to its last remaining
Branchlets slender, flexible, stout and firm on cone refugia in southern Greece. These closely related firs
bearing branches, yellowish grey or grey, nearly will, unsurprisingly, hybridize readily, and the more
smooth, striated, with short, yellowish or brown dominant genome will in due course prevail, oblit-
pubescence; leaf scars broadly elliptic or circular. erating the other species. This process may not have
Vegetative buds ovoid or conical, ca. 6 × 4 mm, not reached its conclusion, and to reflect this history it
resinous or slightly resinous especially on cone bear- seems appropriate to recognize this taxon as of hy-
ing branches; bud scales broadly ovate, obtuse, brown, brid origin in the way we write its name.
persisting several years. Leaves spirally arranged, on
vegetative branches pectinate in two lateral sets, on
Distribution
leading shoots radial, on coning shoots radial and as-
surgent, (1.2–)1.5–3(–3.5) cm long, 1.5–2.6 mm wide, Bulgaria (Rhodope Mts.), Greece (Macedonia to N
(slightly) twisted at base, linear, flattened, grooved Peloponnisos).
and lustrous dark green above, whitish green below; TDWG codes: 13 BUL GRC
apex entire in all leaves. Stomata in two bands sepa-
rated by a midrib below, often also a few stomata on
Ecology
the adaxial (upper) side. Pollen cones lateral, crowd-
ed, 2 cm long, greenish yellow with purple red mi- Abies ×borisii-regis is a montane species occurring in
crosporophylls. Seed cones lateral, erect; peduncles the Balkan mountains at middle elevations between
short, slightly curved; cone shape cylindrical, with 700 m and 1500 m a.s.l. It can grow mixed with Picea
conical or obtuse apex, 10–15(–20) cm long, 3–5(– abies or occur in more or less pure stands; at lower
6) cm wide, yellowish green, sometimes with pur- elevations it can occur in broad-leaved forest.
ple tinge when immature, ripening to light brown
or reddish brown; cone rachis persistent, narrowly
Conservation
conical, dark brown. Seed scales cyathiform, the up-
per ones more cuneate, length × width at mid-cone IUCN: LC
2.5–3 × 2.5–3.5 cm; surface smooth, pulverulent es-
pecially on the exposed part; upper margin entire;
Uses
base long pedicellate. Bracts linear spathulate, with
a long, caudate cusp, 2.5–3.5(–4) cm long, exserted, The timber of this fir is valuable and used for inte-
slightly recurved at maturity. Seeds cuneate, angular, rior construction, mostly as plywood or veneer. It is
7–9 mm long, yellowish brown; seed wings cuneate, similar to European fir (Abies alba) in all its proper-
10–15 mm long, reddish yellow. ties and uses. It has been planted in European arbo-
reta especially in the 19th century and many of these
trees still remain alive; it apparently grows to a bet-
Taxonomic notes
ter shape in the British Isles than A. alba.
This taxon was originally described as a species from
the Rhodope Mountains in Bulgaria and Mt. Olym-
 Abies bracteata (D. Don) A. Poit., Rev. Hort., sér. ish brown. Seed scales ovate or broadly reniform,
2, 4: 7. 1845. Pinus bracteata D. Don, Trans. Linn. length × width at mid-cone 1.5–1.7 × 2–2.2 cm; sur-
Soc. London 17: 443. 1836. Type: USA: California, face smooth or slightly rough, glabrous; upper mar-
Monterey Co., Santa Lucia Mts., D. Douglas s.n. gin entire, incurved; base short pedicellate. Bracts
(holotype K). Pl. 1, Fig. 3 strongly attached to seed scales, obovate-oblong,
with twisted, extremely elongated cusps, (2.5–)3–
5(–6) cm long, exserted, cusps twisted or spreading.
Etymology
Seeds obovate-cuneate, 8–10 mm long, pale brown;
The species epithet refers to the exceptionally long seed wings (oblique) obovate or cuneate, 10 mm
bract tips spreading out from the seed cones. long, pale brown.
66
Vernacular names Distribution
Bristlecone fir, Santa Lucia fir USA: California (Santa Lucia Mts.).
TDWG codes: 76 CAL
Description
Ecology
Trees to 30–40(–50) m tall, d.b.h. to 1–1.5 m; trunk
monopodial, straight, columnar, terete; crown py- This fir species occurs in the Upper Sonoran and
ramidal, regular, with narrowly tapering top. Bark Transition Life Zones of the coastal Santa Lucia
of young trees smooth, grey or whitish, later black- Mountains, in canyon bottoms and on rocky slopes
ish grey to brown, fissured below. Branches of first at elevations between 600 and 900 m a.s.l. It grows
order long, slender, becoming gradually shorter on rocky, clay or loam rich mountain soils, which
towards the top, horizontal, pendulous below; can be very dry in summer. It usually grows within
branches of second order spreading, then pendant. reach of ground or seepage water or near intermit-
Branchlets firm but slender, the lowest long and tent canyon bottom streams. The climate is warm,
pendulous, greenish brown or reddish to purplish with dry summers and moderate, moist winters, the
brown, turning grey green or olive green, smooth or precipitation ranges from 500 to 1300 mm annually.
slightly ridged, glabrous; leaf scars circular. Vegeta- It is a constituent of a mixed forest with Pinus coul-
tive buds fusiform or conical, with acute apex, ca. teri, P. ponderosa, Pseudotsuga menziesii, Quercus
1.2–2 × 0.5–0.8 cm, not resinous; bud scales triangu- spp., Lithocarpus densiflorus, Acer macrophyllum,
lar, obtuse or acute, opening when dry, light brown, and other species.
only the lower ones persistent for a few years. Leaves
spirally arranged, forming two lateral rows, on
Conservation
shaded shoots relatively wide apart, loosely pecti-
nate, curved or straight, on coning shoots slightly Abies bracteata has a limited range, consisting of a
assurgent, 2.8–6 cm long, 2–3.5 mm wide, strongly reasonably large number of small and often quite
twisted at base, linear, narrowed towards both ends, disjunct subpopulations, each containing from a
often curved, flattened, acuminate with callous few score to several hundred trees. No commercial
tips, shallowly grooved above, lustrous dark or grey exploitation threatens this species and most popula-
green above, with two broad, greenish white bands tions are within protected areas or on public lands
separated by a prominent green midrib below. Sto- where the management is aimed at preserving these
mata in two bands below. Pollen cones lateral, pe- remarkable trees. However, they are susceptible to
dunculate, with reflexed involucral scales, yellow forest fires, which in much of central and southern
brown with purplish micro­sporophylls. Seed cones California have increased in intensity and extent due
lateral, erect; peduncles up to 1 cm; cones ovoid- to a complex of largely anthropogenic factors. A hot
globular to ovoid, 4–5 10 cm long, 4–6 cm wide, with forest fire at the wrong time in the wrong location
obtuse apex, greenish with purple tinge when im- could wipe out an entire subpopulation. This species
mature, turning pale purplish brown to brown when is therefore flagged as Near Threatened.
ripe; cone rachis persistent, cylindric conical, black- IUCN: NT
 3

6 67

4 6

2
7

Pl ate 1. Abies bracteata. 1. Habit of tree. 2. Foliage branch. 3. Seed cone; seed scales with bracts. 4. Detail
of buds and leaves. 5. Pollen cone. 6. Seeds. 7. Leaves.
 few on the adaxial (upper) surface near the apex,
Uses
two bands separated by a midrib below. Pollen cones
Santa Lucia fir is not used for timber but it is an at- lateral, crowded on the underside of branches, 1.2–
tractive and unusual species much valued in col- 1.8 cm long, yellow with red microsporophylls. Seed
lections for botanic gardens and arboreta. It was cones lateral, erect; peduncles short; cone shape cy-
successfully introduced and raised by the famous lindrical, 10–16(–20?) cm long, 3.5–5 cm wide; apex
tree nursery of Veitch & Son near Chelsea in Eng- obtuse or papilliform, greenish brown when imma-
land in 1853, but it only became more common in ture, maturing to light brown; cone rachis persistent,
horticulture later in the 20th century when renewed narrowly conical, brown; seed scales cuneate near
seed collecting was undertaken. Trees in cultivation the top, cyathiform at mid-cone, length × width at
68 often grow much faster and taller than in their natu- mid-cone 2.5–3.5 × 2–3 cm; surface smooth, with
ral habitats in the Santa Lucia Mountains. yellowish brown pubescence on the exposed part;
upper margin undulate or entire, convex; base short
pedicellate. Bracts linear-spathulate, with long cusp,
Abies cephalonica Loudon, Gard. Mag. & Reg. 3–4 cm long, exserted and recurved. Seeds narrowly
Rural Domest. Improv. 14: 81. 1838 [Arbor. Frutic. cuneate, 8 × 5 mm, brown; seed wings cuneate or
Brit.]. Abies alba Mill. var. cephalonica (Loudon) oblong, 15–20 × 8–10 mm, lustrous light brown.
Richt., Pl. Europ. 1: 5. 1890. Type: not designated.
Fig. 4
Taxonomic notes
The genetic purity of A. cephalonica is apparently
Etymology
lacking in much of the presumed former range of
The species epithet denotes its locus classicus, the is- this species (Mitsopoulos & Panetsos, 1987). There
land of Cephalonia in Greece. appears to be a north-south cline of character states,
reflecting introgression with A. alba genes with de-
creasing influence on morphological traits. Pure
Vernacular names
A. cephalonica populations occur on Cephalonia
Greek fir, Grecian fir; Kukunaria (Greek) (the locus classicus), Euboea and in the far south
of Peloponnisos, but individual trees with genuine
characters of this species occur mixed with interme-
Description
diate forms well into Macedonia. Variation is con-
Trees to 30–35 m tall, d.b.h. to 1.5–2 m; trunk mo- siderable in much of the area where these hybrid
nopodial, straight, columnar, terete; crown pyrami- forms are supposed to occur. This hybrid has been
dal, old trees flat topped. Bark of young trees greyish, formally named as Abies ×borisii-regis Mattf. and
tinged with pink or brown, smooth, finally in old was originally described from the Rhodope Moun-
trees fissured below. Branches of first order long, tains in Bulgaria. Other botanists described the hy-
spreading horizontally, pendant below; branches of brid as a variety or subspecies of A. alba (A. alba var.
second order spreading or pendant. Branchlets stout, acutifolia Turrill).
shaded ones slender, shining pale brown or reddish
brown, rarely yellowish, prominently grooved, gla-
Distribution
brous; leaf scars circular. Vegetative buds ovoid or
conical, ca. 6 × 4–5 mm, with yellowish resin; bud Greece (Cephalonia, Euboea, Sterea Hellas, Pelo-
scales ovate, acute, reddish brown and persistent ponnisos).
for several years. Leaves spirally arranged, spread- TDWG codes: 13 GRC
ing radially, more or less pectinate below, assurgent
and curved on coning shoots, (1.5–)2–3(–3.5) cm
Ecology
long, 2–2.5 mm wide, twisted or curved at base, lin-
ear, curved and flattened, shining dark green above, Abies cephalonica is a montane species, occurring
with two whitish green stomatal bands separated by from (600–)800–2000(–2100) m a.s.l. in the high
a midrib below; apex acute, rarely obtuse. Stomata mountains of Greece. Soils are usually well drained
and calcareous, but in the north, where introgres- Abies chensiensis Tiegh., Bull. Soc. Bot. France 38:
sion with A. alba seems to have occurred, also on 413. 1892.
­siliceous soils, which may be slightly acid. The cli-
mate has relatively dry summers and wet winters,
Etymology
with annual precipitation from 700 to 1500 mm. At
lower elevations A. cephalonica mixes with Fagus The species epithet refers to the Chinese province
orientalis, Quercus spp., Castanea sativa, and also (now spelled Shaanxi) from where this species was
Pinus nigra, but higher in the mountains it usually first described.
forms pure forests with occasional Juniperus oxy­
cedrus.
Vernacular names
69
Shensi fir; qin ling leng shan (Chinese)
Conservation
Abies cephalonica has a limited distribution, but its
Description
actual extent of occurrence (EOO) is difficult to
establish due to inferred past and perhaps present Trees to 50(–70?) m tall, d.b.h. to 2–2.5 m; trunk
hybridization with A. alba. Where it is considered monopodial, straight, columnar, terete; crown py-
genetically pure, the introduction of firs from other ramidal, becoming flat topped in old trees. Bark of
sources would potentially threaten this species; it young trees dark grey, smooth, fissured in old trees.
is therefore imperative for foresters to refrain from Branches of first order short, massive; branches of
such introductions among or near such popula- second order stout, spreading. Branchlets stout,
tions. The situation becomes more obscure in the firm, yellowish or grey, shining, ridged between
northern Peloponnisos and in Sterea Hellas where leaf scars, glabrous or slightly pubescent at first; leaf
it is found that both A. cephalonica and A. ×borisii- scars roundish with a light centre. Vegetative buds
regis already occur. Climate change may influence conical or ovoid, 10 × 6 mm or larger on some lead-
this situation, with drier conditions presumably ing shoots, (slightly) resinous; bud scales triangu-
benefiting A. ­cephalonica and wetter conditions lar or broadly ovate, brown or reddish, persisting
A. alba, and therefore ongoing introgression of the several years. Leaves spirally arranged, the lower
genes of A. alba into the remaining populations of ranks pectinate at right angles to shoot, the upper
A. ­cephalonica. leaves shorter and directed forward, assurgent or
IUCN: NT nearly erect on coning shoots, (1.3–)2–4.5(–7.5) cm
long, 2.5–3 mm wide, twisted at base, ligulate linear,
flattened, grooved and shining green above, whit-
Uses
ish green below, emarginate or obtuse, occasion-
Greek fir, due to its rarity, is of limited commercial ally acute (coning shoots). Stomata in two broad
value as a timber tree. Crosses with other European bands divided by a midrib below. Pollen cones lat-
firs (A. nordmanniana, A. pinsapo) have been es- eral, crowded, 1 cm long. Seed cones lateral, erect on
tablished and tried or used in plantation forestry short peduncles, oblong-ovoid or cylindrical, with
as well as the natural hybrid A. ×borisii-regis. The a truncate apex, (7–)8–11(–14) cm long, (3–)4–5 cm
latter is more widespread and, together with Euro- wide, green when immature, ripening to cinnamon
pean fir (A. alba), exploited for timber in its natural brown; cone rachis persistent, cylindric-conical,
range. Abies cephalonica was widely planted as an dark brown. Seed scales broadly ovate-cuneate to
ornamental tree in Europe during the 19th century, cyathiform, length × width at mid-cone 2.5–3 ×
distributed from seed collected on Cephalonia. Such 3–3.5 cm; surface smooth, tomentose-puberulent;
trees are now becoming rare because seedlings from upper margin rounded, obscurely serrate; base pedi-
them will not be genetically pure and new collec- cellate. Bracts short, 2-laminated, with short cusp,
tions are seldom made. A few cultivars are known length 1–1.5 cm, entirely included. Seeds obovoid, 10
but these, too, are rare. × 5 mm, brown; seed wings obdeltoid, 20 × 10 mm,
pale brown.
 Distribution Conservation
China: SE Gansu, Henan, W Hubei, Shaanxi, W Si- Logging and deforestation have depleted this most
chuan, NW Yunnan, SE Xizang [Tibet]; NE India: common and widespread subspecies. The assess-
Arunachal Pradesh. ment was based on the 1994 IUCN criteria which
TDWG codes: 36 CHC-HU CHC-SC CHC-YN CHN- have changed in the 2001 IUCN criteria, a new as-
GS CHN-SA CHS-HE CHT 40 EHM-AP sessment is needed.
IUCN: VU (A2d)
Ecology
Abies chensiensis Tiegh. subsp. salouenensis
70 This species occurs in high mountain ranges of the (Bordères & Gaussen) Rushforth, Notes Roy. Bot.
SW Plateau of China between 2100 and 3500 m a.s.l., Gard. Edinburgh 41: 539. 1984. Abies salouenensis
on grey-brown mountain podzols, brown earth or Bordères & Gaussen, Trav. Lab. Forest. Toulouse
lithosols. The climate is cold and moist, with annual T. 1 (4, 15): 4. 1947; Abies ernestii Rehd. var. salou­
precipitation between 1000 and 2000 mm. It is a rare enensis (Bordères & Gaussen) W. C. Cheng &
species, usually mixed with Picea spp., Abies fargesii L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 93. 1978; Abies
var. sutchuenensis, Tsuga chinensis, Larix potaninii at recurvata Mast. var. salouenensis (Bordères &
high elevations, and Betula spp. at lower elevations; Gaussen) C. T. Kuan, Fl. Sichuan. 2: 48. 1983; Abies
also as a pure forest in Tsin-ling Shan (Wang, 1961). chensiensis Tiegh. var. salouenensis (Bordères &
Gaussen) Silba, Phytologia 68: 10. 1990. Type:
China: Yunnan, Hengduan Shan, Atuntze, T. T. Yü,
Uses
7952 (holotype TLF).
This fir has been logged heavily for timber in north-
ern China. As of most tall firs, the wood is soft,
Description
even-grained and suitable for many applications in
construction, veneer, panelling etc. The species is Leaves 4–7.5 cm long; resin canals in leaves of coning
uncommon in cultivation outside China and mostly shoots marginal.
restricted to botanic gardens and arboreta.
Taxonomic notes
Three subspecies are recognized:
In Flora of China 4: 52 (1999) this taxon is classified
Abies chensiensis Tiegh. subsp. chensiensis. Type: as Abies ernestii Rehd. var. salouenensis (Bordères &
China, Shaanxi, Qinling Shandi, Abbé David 918 Gaussen) W. C. Cheng & L. K. Fu; A. ernestii Rehd.
(holotype P). is treated as a variety of A. recurvata in Farjon (1990,
1998, [2001]).
Description
Distribution
Leaves (1.3–)2–4.5(–5) cm long; resin canals in leaves
of coning shoots medial. Seed cones (7–)8–11(–12) SW China: NW Yunnan, SE Xizang [Tibet]; NE In-
cm long. dia: Arunachal Pradesh.
TDWG codes: 36 CHC-YN CHT 40 EHM-AP
Distribution
Conservation
China: SE Gansu, Henan (Nexiang), W Hubei,
S Shaanxi, W Sichuan. IUCN: LC
TDWG codes: 36 CHC-HU CHC-SC CHN-GS CHN-
SA CHS-HE
Abies chensiensis Tiegh. subsp. yulongxueshanensis
Vernacular names
Rushforth, Notes Roy. Bot. Gard. Edinburgh
41: 539. 1984. Abies chensiensis Tiegh. var. Cilician fir
yulongxueshanensis (Rushforth) Silba, Phytologia
68: 10. 1990. Type: China: Yunnan, Lijiang Xian,
Description
Yulongxue Shan, T. T. Yü, 15050 (holotype E).
Trees to 30 m tall, d.b.h. to 0.75–1 m; trunk mo-
nopodial, straight, columnar, terete; crown (nar-
Description
rowly) conical. Bark of young trees grey, smooth, of
Seed cones 10–14 cm long. Resin canals in leaves of old trees scaly and fissured. Branches of first order
coning shoots marginal. spreading horizontally, the upper ones ascending; 71
branches of second order spreading horizontally.
Branchlets stout and firm, yellowish or greenish
Taxonomic notes
brown, turning grey later, shallowly grooved, faintly
Rushforth (1984) described this subspecies from the pubescent or glabrous on first year shoots, all soon
main massif of the Lijiang Shan (Yulongxue Shan, glabrous; leaf scars circular, dark. Vegetative buds
5596 m) in NW Yunnan. It has somewhat larger ovoid conical, acute, small, 3–4 × 2–3 mm, not or
cones than the type subspecies and marginal resin slightly resinous; bud scales broadly ovate, light
canals in the leaves of cone-bearing shoots. In Flora brown or orange brown, persisting several years.
of China 4 (1999) where no subspecies of A. chen- Leaves spirally arranged, directed forward especially
siensis are recognized, this taxon has been instead on vegetative shoots, the lower ranks pectinate, on
treated under Abies ernestii Rehd. var. salouenensis coning shoots radially and assurgent, more or less
(Bordères & Gaussen) W. C. Cheng & L. K. Fu as a forward, 2.5–4 cm long, 1.5–1.8 mm wide, twisted
synonym. and abruptly narrowed at base, linear, flattened,
bright green or glaucous above, greenish white with
green midrib below; apex obtuse or slightly emar-
Distribution
ginate. Stomata in two broad bands separated by a
China: Yunnan (Lijiang Shan). midrib below. Pollen cones lateral, axillary, short
TDWG codes: 36 CHC-YN pedunculate, 1–1.5 cm long, yellowish with red mi-
crosporophylls. Seed cones lateral, erect, often in
pairs, with short peduncles or sessile, cylindrical,
Conservation
with obtuse, retuse or papilliform apex, 16–20(–30)
This subspecies is only known from one mountain cm long, 4–6 cm wide, greenish when immature,
range in NW Yunnan and is believed to count fewer turning reddish brown (often resinous) and brown
than 1000 mature trees. Parts of this mountain are with age; cone rachis persistent, narrowly conical,
a National Park and at least some of the population dark brown. Seed scales broadly flabellate, length ×
occurs within this protected area. width at mid-cone 2.5–3 × 3–4 cm; surface smooth,
IUCN: VU (D2) pubescent on exposed parts; upper margin thin and
entire; base pedicellate. Bracts narrowly spathulate,
1.5–2.5 cm long, usually included, sometimes slightly
Abies cilicica (Antoine & Kotschy) Carrière, Traité exserted. Seeds obovate-triangular, 10–12 × 5 mm,
Gén. Conif.: 229. 1855. light brown; seed wings cuneate, broad, 14–17 mm
long, light brown.
Etymology
Taxonomic notes
The species epithet means ‘from Cilicia’ the classi-
cal name for a part of the Mediterranean coast of The minor and probably variable differences be-
Turkey. tween A. cilicica subsp. cilicica and subsp. isaurica
are considered by some specialists in the genus to be
 insignificant, but because the latter subspecies was
Distribution
recognized in Vol. 1 of the Flora of Turkey (Davis et
al., 1965), and as it occurs only in the Isaurian Taurus Lebanon, Syria, Turkey (Anti-Taurus, Cilician Tau-
Mountains, it is here separated from subsp. cilicica. rus).
TDWG codes: 34 LBS-LB LBS-SY TUR
Distribution
Conservation
Lebanon, Syria, Turkey.
TDWG codes: 34 LBS-LB LBS-SY TUR IUCN: LC

72 Abies cilicica (Antoine & Kotschy) Carrière subsp.

Ecology
isaurica Coode & Cullen, Notes Roy. Bot. Gard.
Abies cilicica is a species of the high mountains Edinburgh 26: 167. 1965. Type: Turkey: Antalya,
around the eastern Mediterranean Basin, occurring Gebiz, Bozburun Dag, [“Bozburun Dag, between
at elevations between 1200 m to 2100 m a.s.l. (subsp. Bogarz Azzi & Tozlu Cukur yoyla”(?)], P. H. Davis
cilicica) and 1000 m to 2000 m a.s.l. (bsp. isaurica). 15505 (holotype E).
The soils are often shallow and rocky, well drained,
calcareous, and dry in the summer. The climate is
Description
characterized by warm, dry summers and wet, mild
winters, the annual precipitation is 1000 to 1500 New shoots glabrous; buds slightly resinous.
mm. In its lowest reaches Quercus spp. and Taxus
baccata are mixed in, but higher on the mountains
Distribution
it forms pure stands or mixed forests with Cedrus
libani; Pinus nigra and Juniperus excelsa occur more Turkey: Prov. Antalya and Konya (Isaurian Taurus).
rarely. TDWG codes: 34 TUR

Uses Conservation
Cilician fir is an important timber tree in Turkey, This subspecies (if distinct) has a limited distribu-
where it is relatively abundant. Its wood is used for tion and therefore, dependent on trends in its abun-
indoor construction mainly as plywood. It is rare dancy which are currently unknown, it could in the
in cultivation despite its alleged horticultural mer- near future qualify for a category of threat. The re-
its because of its susceptibility to ‘late’ frost damage gion has experienced substantial forest depletions in
which can occur in much of western Europe due to the past.
erratic cold and warm spells in early spring. IUCN: NT

Two subspecies are recognized:

Abies concolor (Gordon) Lindl. ex Hildebr., Verh.
Abies cilicica (Antoine & Kotschy) Carrière subsp. Naturhist. Vereines Preuss. Rheinl. Westphalens 18:
cilicica. Pinus cilicica Antoine & Kotschy, Oesterr. 261. 1861. Picea concolor Gordon, Pinetum: 155. 1858.
Bot. Wochenbl. 3: 409. 1853. Type: Turkey: Bulgar Type: USA: New Mexico, A. Fendler 828 (holotype
Daglari, K. G. T. Kotschy 49611 (holotype G). MO).

Abies cilicica (Antoine & Kotschy) Carrière var. Picea lowiana Gordon, Pinetum Suppl.: 53. 1862;
p­ yramidalis Boydak & Erdogrul, Istanbul Univ. Or- Abies lowiana (Gordon) A. Murray bis, Proc. Roy.
man Fak. Dergisi, A, 49 (2): 21. [1999] 2000. Hort. Soc. London 1863 (3): 317. 1863; Abies grandis
(Douglas ex D. Don) Lindl. var. lowiana (Gordon)
Hoopes, Book Evergr.: 212. 1868; Abies concolor
Description
(Gordon) Lindl. ex Hildebr. var. lowiana (Gordon)
New shoots faintly pubescent, soon glabrous; buds Lemmon, Cone-Bear. Trees Pacif. Slope, ed. 3: 64.
not resinous. 1895; Abies concolor (Gordon) Lindl. ex Hildebr.
subsp. lowiana (Gordon) E. Murray, Kalmia 13: 3. apex, 7–12(–15) cm long, 3–4.5 cm wide, colour vari-
1983. able when immature, usually light (yellowish )green,
Abies concolor (Gordon) Lindl. ex Hildebr. f. atro- sometimes purple violet, maturing to (light) brown;
violacea Cinovskis, Introd. Rast. Bot. Sad. Pribaltiki: cone rachis persistent, narrowly conical, black-
48. 1974. ish brown. Seed scales flabellate, sometimes much
Abies concolor (Gordon) Lindl. ex Hildebr. var. baja- wider than long, length × width at mid-cone 2.5–3
californica Silba, Phytologia 68: 11. 1990. × 3–3.5 cm; surface smooth, puberulent, especially
Abies concolor (Gordon) Lindl. ex Hildebr. var. mar- on exposed parts; upper margin entire; base pedi-
tinezii Silba, Phytologia 68: 12. 1990. cellate, often curved. Bracts ovoid-oblong, with two
Abies lowiana (Gordon) A. Murray bis var. viridula lateral wings and short cusp, 1–1.5 cm long, includ-
Debreczy & Rácz, Phytologia 78 (3): 13. 1995. ed. Seeds cuneate, 8–10 mm long, dull light brown; 73
seed wings oblong-cuneate, 15–20 mm long, light,
lustrous brown.
Etymology
The species epithet refers to the similar colour on
Taxonomic notes
both sides of the leaves.
Investigation of herbarium specimens from loca-
tions covering much of the range of A. concolor
Vernacular names
revealed a great variety in arrangement and length
White fir, Colorado fir, Low’s fir, California white fir of leaves. It has been concluded that var. lowiana
(Gord.) Lemmon falls entirely within the variation
of var. concolor as found in specimens from Rocky
Description
Mountain locations (Farjon, 1988). There is conse-
Trees to 60–70 m tall, d.b.h. to 2–2.5 m; trunk mo- quently no taxonomically sound basis upon which
nopodial, straight, columnar, terete; crown nar- to recognize this perceived entity as a distinct spe-
rowly conical, with flat domed top in old trees. cies or as an infraspecific taxon.
Bark of young trees smooth, grey to whitish with
conspicuous resin blisters; in old trees dark grey,
Distribution
tinged with yellowish brown, scaly, deeply fissured
below. Branches of first order relatively short, in USA: from Oregon and Idaho through the Rocky
regular, horizontal whorls in young trees, the lower Mountains and Californian mountains to Arizona.
ones later drooping, curved; branches of second or- Mexico: Baja California Norte, Chihuahua, Sonora.
der spreading. Branchlets slender, yellowish green, TDWG codes: 73 COL IDA ORE WYO 76 ARI CAL
soon becoming grey brown, smooth or with shal- NEV UTA 79 MXE-CU MXN-BC MXN-SO
low grooves, glabrous or with minute pubescence
on first year shoots; leaf scars circular, light. Vegeta-
Ecology
tive buds globose or globular, 3–5 mm long, resin-
ous; bud scales ovate triangular, brown, persistent Abies concolor occurs in the Transition and Cana-
for 2–3 years. Leaves spirally arranged, in irregular dian Life Zones of high mountains, often restricted
ranks, spreading, more or less pectinate in some to N-facing slopes, at elevations between 600 to
trees, curved upward and forward in others, assur- 3000 m in the western part of its range, from 1800
gent or nearly erect on coning shoots, (1.5–)4–6(–7) m to 3350 m a.s.l. in the Rocky Mountains. It grows
cm long, 2–3 mm wide, curved or (strongly) twisted in a variety of mountain soils on granitic or basal-
at base, linear, usually curved, or the shortest leaves tic rocks or occasionally sandstone. The climate is
falcate, flattened or slightly carinate, green or glau- moderately humid (500 to 1875 mm annual precipi-
cous green; apex obtuse or acute. Stomata in a few tation), with relatively warm and dry summers and
or more lines above, two bands divided by a midrib cold winters. It grows in pure stands, or mixed with
below. Pollen cones in the axils of leaves, pendulous, various other conifers, e.g. Pinus spp., Abies magnifi-
1.2–2 cm long, with red or roseate microsporophylls. ca, A. procera, A. grandis, Pseudotsuga menziesii var.
Seed cones lateral, erect, sessile or very short pedun- glauca, and also with Populus tremuloides.
culate, cylindrical or elliptical, with obtuse or retuse
 arranged, in several radially spreading, overlapping
Conservation
ranks of different length, more or less forward, on
IUCN: LC coning shoots more assurgent, (1.2–)1.5–3(–4.3) cm
long, 1–2(–2.8) mm wide, twisted at base, linear, of-
ten curved or S-shaped, above lustrous dark green,
Uses
below niveous white, often also with white wax on
White fir provides timber for general construction the midrib; margins strongly revolute; apex emar-
applications and plywood; its market value has im- ginate or obtuse; stomata in two bands below. Pollen
proved in recent decades relative to other species of cones laterally on the lower side of branches, pen-
fir. In California it is grown for Christmas trees in dant from the leaf axils, 2–3.5(–4) cm long, yellow
74 plantations or harvested from natural regeneration with violet microsporophylls. Seed cones lateral,
in forests managed for timber production. The natu- erect; peduncles short, with many scales; shape cy-
ral variation in leaf colour from green (recognized lindrical or ovoid-oblong, with truncate or slightly
as var. lowiana in horticulture) to glaucous green umbilicate apex, 6–11(–14) cm long, 3–4.5 cm wide,
can be enhanced by selection and has been used dark violet blue when immature, turning purplish
both in the Christmas tree trade and in horticulture. black or blackish brown with blue bracts, dull (red-
A number of cultivars, including dwarfed forms, are dish) brown when ripe; cone rachis persistent, thick,
commonly used in gardens. conical to fusiform, purplish brown. Seed scales
­obtriangular-flabellate to flabellate-reniform, auric-
ulate, length × width at mid-cone 1.2–2 × 1.5–2.5 cm;
Abies delavayi Franch., J. Bot. (Morot) 13 (8): 255. surface smooth, puberulent; apical part thickened,
1899. with rounded, slightly recurved, entire margin, lat-
eral margins erose; base pedicellate. Bracts oblong-
spathulate, straight or recurved at apex, with small
Etymology
or long, narrow cusps, 1–2 cm long, entirely includ-
This species was named after the French mission- ed or exserted, usually with spreading cusps. Seeds
ary P. J. M. Delavay, who collected plants in Yunnan, obovate, 5–8 mm long, brown (mostly covered with
China, during the last quarter of the 19th century. a blackish wing membrane); seed wings cuneate-
dolabriform, 10 × 6 mm, (light) brown with a black
or purplish blue tinge.
Vernacular names
Delavay’s fir; cang shan leng shan (Chinese)
Distribution
SW China: W Yunnan, SE Xizang [Tibet]; NE India:
Description
Arunachal Pradesh; N Myanmar [Burma]; N Viet-
Trees to 30–40 m tall, d.b.h. to 1–1.5 m; trunk mo- nam.
nopodial, straight, columnar, terete; crown (broad) TDWG codes: 36 CHC-YN CHT 40 EHM-AP 41 MYA
pyramidal, flat topped in old trees. Bark of young vie
trees grey or brownish grey, smooth, in old trees
deeply fissured and flaking off in small fragments.
Ecology
Branches of first order massive, spreading, ascend-
ing near the top; branches of second order spreading, This is a species of high elevations in the great
assurgent. Branchlets stiff, terete, maroon or brown, mountain ranges of SW China, occurring between
usually dull, sometimes shining dark red brown or 2400 m and 4300 m a.s.l., but usually between 3000
purplish brown, becoming grey, smooth or slightly m and 4000 m, commonly on N-facing slopes. The
grooved, glabrous, puberulent or ferruginous pu- soil is a grey brown mountain podzol. The climate is
bescent in grooves on 1st year shoots; leaf scars angu- extremely wet, with cool summers and cold, snowy
lar circular. Vegetative buds ovoid-globular, small, winters (annual precipitation ranges from 1000 mm
or up to 8 mm long on leading shoots, very resinous; to 3000 mm and more). It grows mixed with other
bud scales broadly ovate, obtuse, reddish or green- conifers, such as Picea likiangensis, P. brachytyla var.
ish brown, persisting several years. Leaves spirally brachytyla, or in pure stands towards the tree lim-
it. At lower elevations it is sometimes mixed with Abies delavayi Franch. subsp. fansipanensis
Tsuga chinensis, T. dumosa, Juniperus formosana and (Q. P. Xiang) Rushforth, Int. Dendrol. Yearb. 1998:
broad-leaved trees, e.g. Betula albosinensis, Betula 62. 1999. Abies fansipanensis Q. P. Xiang, Acta
platyphylla var. szechuanica, and Quercus semecarpi- Phytotax. Sin. 35 (4): 356. 1997. Type: Vietnam, Lao
folia. Abies delavayi, however, is less common with Cai Prov., Mt. Fan Si Pan, Sino-Vietnamese Exped.
these trees than A. forrestii. s.n. (holotype PE)

Uses Description
This high altitude species is not an important timber Trees 15–20 m tall. Seed cones cylindrical, 8–14 cm
tree, but it has been logged in several areas together long; bracts bilobed, with small cusp, entirely in- 75
with other conifers with which it occurred. Measures cluded.
to reduce or even stop logging these high mountain
forests are being put in place by the Chinese govern-
Distribution
ment. This species, and some of its forms or varie-
ties, have been introduced to cultivation in Europe, Vietnam: Lao Cai Prov. (Mt. Fan Si Pan).
in particular the UK and France, but are still mostly TDWG codes: 41 VIE
restricted to arboreta and botanic gardens, as they
are slow growing.
Ecology
Three varieties and one subspecies are recognized: Abies delavayi subsp. fansipanensis is endemic on
Mt. Fan Si Pan, the highest mountain in Vietnam
Abies delavayi Franch. var. delavayi. Type: China, in the extreme NW of the country. It is an outlier of
Yunnan, Tsang Shan, near Dali, P. J. M. Delavay 1210 a widespread species with a typical Sino-Himalay-
(holotype P). Fig. 5, 6 an distribution. It grows on the highest slopes and
ridges of this granitic mountain at altitudes between
2600 m and 3000 m a.s.l. as an emergent tree in pri-
Description
mary evergreen tropical high montane forest. Rain-
Shoots glabrous or puberulent in grooves; leaves fall is very high, to 3,500 mm or more per year and
(1.2)1.5–3(–3.5) cm long, 1–2 mm wide, strongly clouds cover the summit region of the mountain for
revolute. Seed cones ovoid-oblong to cylindrical, most of the time. It is associated with Tsuga dumosa
6–11(–14) cm long; bracts (slightly) exserted, with a (which has a similar geographical distribution) and
long, narrow cusp. Rhododendron spp.; the understorey is often domi-
nated by bamboos.
Distribution
Conservation
SW China: W Yunnan, SE Xizang [Tibet]; NE India:
Arunachal Pradesh; N Myanmar [Burma]. At present, there are no direct threats from log-
TDWG codes: 36 CHC-YN CHT 40 EHM-AP 41 MYA ging due to its remote habitat, but if plans to build a
road on the mountain were to materialize this may
change, despite the fact that the entire population
Conservation
occurs within the Hoang Lien National Park. In-
IUCN: LC creasing tourism on the mountain could heighten
the risk of fires in dry spells.
IUCN: VU (D)
 Abies delavayi Franch. var. motuoensis W. C. Cheng
Etymology
& L. K. Fu, Acta Phytotax. Sin. 13 (4): 83. 1975. Type:
China, SE Xizang [Tibet], Motuo, Chinese collector The species epithet refers to the crowded leaves.
1011 (holotype PE)
Vernacular names
Description
Sikkim fir; gobria, gobra salla (Nepal); dunshing
Shoots densely ferruginous pubescent; leaves 2–3.2 (Bhutan)
cm long.
Description
76
Distribution
Trees to 50–60 m tall, d.b.h. to 2.5 m; trunk mo-
China: SE Xizang [Tibet] (Motuo). nopodial, straight, columnar, terete; crown broad,
TDWG codes: 36 CHT pyramidal or columnar, in old trees flat topped.
Bark soon scaly, grey, in old trees fissured towards
base, breaking into large plates. Branches of first or-
Conservation
der thick, long, assurgent near the top, horizontally
IUCN: LC spreading, or lower branches bent down; branches
of second order assurgent. Branchlets thick in lead-
Abies delavayi Franch. var. nukiangensis ing shoots, lateral shoots firm but slender, yellow-
(W. C. Cheng & L. K. Fu) Farjon & Silba, Phyto- ish to reddish brown, becoming grey, prominently
logia 68: 13. 1990. Abies nukiangensis W. C. Cheng ridged and grooved, glabrous or with slight pubes-
& L. K. Fu, Acta Phytotax. Sin. 13 (4): 83. 1975. cence in grooves; leaf scars circular ovate. Vegeta-
Type: China: Yunnan, Hengduan Shan, Nu Jiang, tive buds ovoid-conical, 8 × 6 mm, resinous; bud
K. M. Feng 8025 (holotype PE). Fig. 7 scales ovate, obtuse, orange brown and persistent
for several years. Leaves spirally arranged, more or
less pectinate, on leading and coning shoots more
Description
radially arranged, with the upper leaves curved
Shoots glabrous, purplish brown; leaves 1.2–4.3 cm forward, (1.5–)2–4(–5) cm long, 1.5–2.5 mm wide,
long, 1.5–2.8 mm wide, sometimes only slightly revo- twisted or curved at base, linear, straight or slightly
lute. Cone bracts with a short cusp, mostly included. curved, flattened; margins slightly revolute; upper
surface of leaves light or mid green, lower surface
with two white bands; apex emarginate. Stomata in
Distribution
two bands separated by a midrib below. Pollen cones
China: NW Yunnan (Nukiang River). lateral, radially arranged around the shoot, 2–4.5 cm
TDWG codes: 36 CHC-YN long, yellow with purplish blue microsporophylls.
Seed cones lateral, erect, sessile or on very short
peduncles, (ovoid-)cylindrical with obtuse or um-
Conservation
bilicate apex, 8–12 cm long, 4–5.5 cm wide, purplish
IUCN: LC blue when immature, becoming blackish purple
with some brown when ripe. Seed scales flabellate-
cuneate, length × width at mid-cone 1.5–2 × 2–2.5
Abies densa Griff., Notul. Pl. Asiat. 4: 19. 1854. Abies cm; surface smooth or slightly striated, pubescent
spectabilis (D. Don) Spach var. densa (Griff.) Silba, on exposed parts; upper margin entire, incurved;
Phytologia Mem. 7: 10. 1984. Type: not designated. base pedicellate. Bracts ligulate-spathulate, 2–2.5 cm
long, slightly or not exserted. Seeds cuneate, 8 × 4
Abies fordei Rushforth, Int. Dendrol. Soc. Yearb. mm, brown; seed wings cuneate, 10 × 5 mm, brown.
2008: 42. 2009.
 Taxonomic notes Conservation
This species has been included with A. spectabi- IUCN: LC
lis (Liu, 1971; Cheng & Fu, 1978; Sahni [in part],
1990), but it is distinct both morphologically and
Uses
geographically. It has somewhat smaller cones with
slightly exserting bracts and leaves with slightly Sikkim fir is a timber tree of importance in the
revolute margins, which are arranged more radially eastern Himalaya, where it is used in construction
around the shoots, especially on coning branches. In (house building), in particular for interior work
these characters it is intermediate between A. spect- such as floor boards, ceilings and stairs, while shin-
abilis and A. delavayi, the latter a species occurring gles are used for roofing. This species has been in- 77
from NE India to Yunnan, China, and also disjunct troduced in Europe only relatively recently from
in N Vietnam. Rushforth (2009) described a new Bhutan, Nepal and Sikkim. It is still uncommon in
species, A. fordei, from S Xizang [Tibet] ocurring gardens and parks but has more attractive characters
on the northern side of the Himalayan crest in the than A. spectabilis, with very white leaf undersides
Yarlung Zangbo drainage. Apart from its flat, not and deep purplish blue, nearly black seed cones. It
revolute, leaf margins, it is similar to A. densa and undoubtedly requires ample precipitation, judging
is here treated as synonymous. It thus appears that from its natural habitat.
at high elevation in the Himalaya, we find from west
to east, with partly overlapping distributions, first
A. spectabilis, then A. densa, and finally A. delavayi, Abies durangensis Martínez, Anales Inst. Biol.
which extends into the adjacent mountains of west- Univ. Nac. México 13 (2): 621. 1942.
ern China and as far as Vietnam.
Etymology
Distribution
The species epithet refers to the Mexican state of
E Himalaya: from E Nepal to the Great Bend of the Durango.
Brahmaputra.
TDWG codes: 36 CHT 40 ASS-AS EHM-AP EHM-BH
Vernacular names
EHM-DJ EHM-SI NEP
Durango fir
Ecology
Description
Abies densa occurs in the high mountains of the
eastern Himalaya, from 2450 m to 4000 m a.s.l., on Trees to 30–40 m tall, d.b.h. to 1–1.5 m; trunk mo-
rocky, often steep slopes in the cloud belt, where it nopodial, straight, columnar, terete; crown narrowly
grows on a variety of alpine lithosols. The climate pyramidal or more rounded, lower half of trunk of-
is extremely wet, with well marked monsoons and ten bare of branches. Bark of young trees smooth,
an annual precipitation of more than 2000 mm. The greyish or reddish brown, in old trees dark blackish
summers are relatively warm, the winters are cold brown and deeply fissured below. Branches of first
at high altitudes and bring heavy snowfall. The spe- order long, spreading horizontally; branches of sec-
cies occurs in a wide altitudinal range from mixed ond order spreading or ascending. Branchlets firm,
deciduous coniferous forest at lower elevations to slender, dark purplish red or red-brown, turning
stands with Betula utilis at tree line. Deciduous trees brown with age, prominently ridged and grooved
are e.g. Acer caudatum, A. pectinatum, Prunus spp., between the leaves, glabrous or sparsely to densely
Sorbus spp. and many large Rhododendron spp. Most pubescent in grooves; leaf scars circular or somewhat
of these disappear above 3000 m to make place for ovate. Vegetative buds oblong-ovoid, 4–5 mm long,
conifers. Picea spinulosa and Tsuga dumosa occur very resinous; bud scales dark brown (but covered
generally in a belt below Abies; Larix griffithiana with yellowish resin), persisting 2–3 years. Leaves
and/or Juniperus squamata above it, the latter at tree spirally arranged, more or less pectinate, curved
line. slightly backward or forward, on coning shoots
 the same or weakly assurgent, (1.4–)2–3.5(–4.5) from high altitude provenances are being grown
cm long, 1–1.6 mm wide, twisted at base, narrowly with some success, but mostly slowly, in the south of
linear, tapering towards an acutish or obtuse apex, England and in Belgium.
(dark) green or light green; none or with a few lines
of stomata above, two bands separated by a midrib 2 varieties are recognized:
below. Pollen cones lateral, in leaf axils, 1–2 (?) cm
long (according to Liu, 1971, 5 mm, but likely longer Abies durangensis Martínez var. durangensis.
when mature), with resinous perular scales; micro- Type: Mexico, Chihuahua, Pinos Altos, Dec 1939,
sporophylls red. Seed cones lateral, erect, on very J. H. Faull s.n. (holotype MEXU)
short peduncles, cylindrical with obtuse apex, 5.5–10
78 cm long, 3–4.5 cm wide; pale brown or ochraceous Abies neodurangensis Debreczy, Rácz & Salazar,
when immature, ripening to pale brown; cone rachis Phytologia 78 (3): 7. 1995.
persistent, narrowly conical. Seed scales flabellate-
cuneate, usually wider than long, length × width at
Description
mid-cone 1.5–2 × 2–2.8 cm; surface smooth, pubes-
cent on exposed parts; upper margin entire, slightly Shoots glabrous or sparsely pubescent in grooves;
reflexed; base pedicellate. Bracts spathulate, with leaves 2–3.5(–4.5) cm long, with bands of stomata
short cusps, 1–1.5 cm long, included. Seeds cuneate- wider than the midrib, usually a few lines of stomata
oblong, 7–8 mm long, yellowish brown; seed wings on the upperside.
cuneate to more or less rounded, 8–10 × 6–8 mm,
pale yellowish. Distribution
Mexico: Chihuahua, Durango, N Jalisco, Sinaloa.
Distribution TDWG codes: 79 MXE-CU MXE-DU MXN-SI MXS-JA
Mexico: Chihuahua, Coahuila, Durango, N Jalisco,
Conservation
Sinaloa.
TDWG codes: 79 MXE-CO MXE-CU MXE-DU MXN- IUCN: LC
SI MXS-JA
Abies durangensis Martínez var. coahuilensis
(I. M. Johnst.) Martínez, Pinaceas Méxic., ed. 3:
Ecology
139. 1963. Abies coahuilensis I. M. Johnst., J. Arnold
Abies durangensis occurs in high mountain val- Arbor. 24: 332. 1943. Type: Mexico: Coahuila, Char­
leys and steep canyons on the western side of the retera Canyon, I. M. Johnston 9010 (holotype A).
divide of the Sierra Madre Occidental, at elevations
between (1600–)2000–2900 m a.s.l. and usually on
Description
well drained talus or lithosols. The climate is moist
and cool, especially on N-facing slopes. It is a rela- Shoots densely pubescent; leaves 1.4–3 cm long, with
tively rare constituent of coniferous forests in the bands of stomata narrower or as wide as the midrib,
Canadian Life Zone, with Pseudotsuga menziesii few or no stomata on the upperside.
var. glauca, Pinus strobiformis, P. leiophylla var. chi-
huahuana, Cupressus lusitanica, C. arizonica in the
Distribution
north of its range, Picea chihuahuana (only near El
Salto, Durango), Pinus durangensis (in the south), Mexico: Coahuila.
Juniperus dep­peana (locally) and some broad-leaved TDWG codes: 79 MXE-CO
trees like Quercus castanea, Q. rugosa, Prunus se-
rotina.
Ecology
Var. coahuilensis has only been found in steep can-
Uses
yons at 2200 m to 2300 m a.s.l., in dense forest with
Durango fir is not a commercially important tim- Pseudotsuga menziesii var. glauca, Pinus strobiformis
ber tree and is extremely rare in cultivation. Trees and Cupressus arizonica.
 on exposed parts, apical end thickened; upper mar-
Conservation
gin entire, incurved; base pedicellate. Bracts oblong-
IUCN: VU (D2) spathulate, 1–1.5 cm long, only the cusps exserted
and recurved or reflexed. Seeds oblong-ovate, 8 ×
4 mm, light brown (partly covered by the blackish
Abies fabri (Mast.) Craib, Notes Roy. Bot. Gard. wing membrane); seed wings cuneate-dolabriform,
Edinburgh 11: 278. 1919. 10 × 6 mm, blackish brown.

Etymology Distribution
This species was named after Ernst Faber (1839–99), China: W Sichuan. 79
who collected plants in China, among which the TDWG codes: 36 CHC-SC
type specimen of this species.
Ecology
Vernacular names
The type location of Abies fabri is on Mt. Emei (Emei
Faber’s fir; leng shan, pao shan (Chinese) Shan), a mountain SW of Chengdu in Sichuan. The
species occurs there at elevations between 2000 m
and 3100 m a.s.l. [K. D. Rushforth, pers. comm.;
Description
Craib (1919) has given a range between 3000 m and
Trees to 40 m tall, d.b.h. to 1–1.5 m; trunk mono­ 3600 m a.s.l.] in a humid, cool climate (mean temp.
podial, straight, columnar, terete; crown often open, in Jan. –4 °C, in July +12.6 °C, annual precipitation
broad pyramidal or flat topped in old trees. Bark >2000 mm). There are some nearly pure stands
of young trees smooth, grey, deeply fissured below and scattered trees on Mt. Emei, but elsewhere in
in old trees. Branches of first order long, spreading W ­Sichuan the species occurs mixed with Picea
horizontally, ascending near the top; branches of ­likiangensis, Tsuga chinensis and occasionally Larix
second order spreading and assurgent. Branchlets potaninii.
stiff, terete, shiny, light yellow or brown, grooved
and ridged between the leaves, glabrous or with a
Uses
few dark hairs in the grooves; leaf scars circular.
Vegetative buds ovoid-conical, small, resinous; bud This species is not known to be a commercially im-
scales ovate, with obtuse apex, greenish purple, portant timber tree, presumably due to its restrict-
persisting several years. Leaves spirally arranged, ed occurrence (protected from exploitation on the
spreading sideways, pectinate below, parted above, ‘holy’ mountain Emei Shan). It is uncommon in cul-
on coning shoots more radially, 1.5–3(–4) cm long, tivation and mostly restricted to arboreta and bo-
2–2.5 mm wide, twisted or curved at base, linear, tanic gardens.
straight or slightly curved, flattened, with revolute
margins; green above, two white stomatal bands Two subspecies are recognized:
separated by a midrib below; apex emarginate or bi-
fid (obtuse or acute on coning shoots). Stomata in Abies fabri (Mast.) Craib subsp. fabri. Keteleeria
two bands separated by a midrib below. Pollen cones fabri Mast., J. Linn. Soc., Bot. 26: 555. 1902; Abies
lateral, axillary, pendant, 2–3.5 cm long, yellow with delavayi Franch. var. fabri (Mast.) D. R. Hunt,
violet or purplish blue microsporophylls. Seed cones J. Roy. Hort. Soc. London 92: 263. 1967. Type:
lateral, erect; peduncles 0.5 cm; shape ovoid-oblong China: Sichuan, Emei Shan, E. Faber 984 (holotype
or more barrel-shaped, with obtuse or umbilicate K).
apex, 5–10 cm long, 3.5–5 cm wide, purplish blue
when immature, ripening to brownish blue; cone
Description
rachis persistent, fusiform, purplish brown. Seed
scales flabellate, auriculate, length × width at mid- Shoots light brown, with a few dark hairs in the
cone 1.4–1.8 × 1.6–2 cm; surface smooth, puberulent grooves or glabrous; leaves 1.5–3 cm long.
 Distribution Vernacular names
China: W Sichuan. fan jing lengshan (Chinese)
TDWG codes: 36 CHC-SC
Description
Conservation
Trees to 22 m tall (or more), d.b.h. to 0.65 m (or
IUCN: LC more); trunk monopodial, straight, columnar, terete;
crown (narrowly) pyramidal or oblong cylindrical.
Abies fabri (Mast.) Craib subsp. minensis (Bordères Bark of young trees grey, smooth, becoming dark
80 & Gaussen) Rushforth, Notes Roy. Bot. Gard. Edin­ grey or greyish brown, scaly and fissured on lower
burgh 43: 273. 1986. Abies minensis Bordères & part of trunk. Branches of first order large, massive,
Gaussen, Trav. Lab. Forest. Toulouse T. 1 (4, 15): spreading horizontally; branches of second order
10. 1947. Type: China: Sichuan, Min River, Songpan, spreading or ascending. Branchlets stout, firm, light
T. T. Yü 8586 (holotype TLF). or dark reddish brown, in 2–3 years becoming dark
brown, ridged and grooved, glabrous or with pubes-
cence in the grooves; leaf scars circular. Vegetative
Description
buds ovoid-globular, 5 × 4 mm, resinous; bud scales
Shoots yellowish, glabrous; leaves (1.8–)3–4 cm long. broadly triangular ovate, reddish brown, persisting
several years. Leaves spirally arranged, crowded, the
lower more or less pectinate, in two lateral rows, the
Taxonomic notes
others more radially spreading or assurgent and di-
In Flora of China 4: 47 (1999) this taxon is treated as rected obliquely forward, often recurved near shoot
a synonym of Abies fargesii var. faxoniana. Its new apex, 1–4.3 cm long (but mostly less than 2.5 cm),
shoots are there described as light brown or grey- 2–3 mm wide, twisted or curved at base, linear, or
brown and being densely rusty brown pubescent on the shorter leaves ligulate linear, flattened, longitu-
lateral branchlets, but usually glabrous on leading dinally grooved and light green above, two whit-
branches. ish bands below; apex (slightly) emarginate, also
of leaves on coning shoots. Stomata in two bands
separated by a narrow midrib below. Pollen cones
Distribution
lateral, axillary, crowded, oblong, 1–1.5 cm long, yel-
China: W Sichuan low, with red microsporophylls. Seed cones lateral,
TDWG codes: 36 CHC-SC erect; peduncles 0.8–1 cm; oval cylindric or barrel
shaped, with truncate or umbilicate apex, 5–6(–7)
cm long, 3.5–4 cm wide, bluish purple when imma-
Conservation
ture, maturing to dark purplish brown, becoming
IUCN: LC dark brown when ripe; cone rachis persistent, fusi-
form, blackish brown. Seed scales broadly cuneate
or reniform, length × width at mid-cone 1.5 × 1.8–
Abies fanjingshanensis W. L. Huang, Acta Phytotax. 2.2 cm; surface smooth, or slightly striated, exposed
Sin. 22 (2): 154. 1984. Abies fargesii Franch. var. part densely puberulent, apically thickened; upper
fanjingshanensis (W. L. Huang) Silba, Phytologia 68: margin entire, rounded, incurved; base pedicellate
15. 1990. Type: China: Guizhou, Yinjiang Tujiazu (‘pedicels’ 8–10 mm long, curved). Bracts spathu-
Miaozu Zizhix, Fanjing Shan, L. Yang 83–427 (holo­ late-obcordate, with a tiny cusp; upper margin
type Herb. Dept. Geogr. Guiyang Teach. Coll., iso­ with irregular fine teeth, 1–1.2 cm long, included or
type PE). slightly exserted. Seeds ovoid-conical, slightly flat-
tened, length × width 6–8 × 3–4 mm, dark brown;
seed wings trapezoid-cuneate or dolabriform, 6–8 ×
Etymology
4–6 mm, light brown or reddish brown.
The species epithet refers to Mt. Fanjing (Fanjing
Shan) on which it occurs.
 Taxonomic notes Vernacular names
This species is closely related to Abies fargesii and is Farges fir; Bashan lengshan (Chinese)
possibly only a subspecies, even though it occurs at
least 500 km to the south of the nearest occurrence
Description
of A. fargesii in the NW of Hubei. Both Farjon (1990)
and Flora of China 4: 46 (1999) have recognized it as Trees to 40 m tall, d.b.h. to 1.5–2 m; trunk monopo-
a distinct species. dial, straight, columnar, terete; crown narrowly
pyramidal or conical. Bark of young trees smooth,
finely flaking, grey, of old trees fissured, brown
Distribution
grey. Branches of first order massive, short, sparse; 81
China: NE Guizhou (Fanjing Shan, near Jiangkou). branches of second order spreading, assurgent, or
TDWG codes: 36 CHC-GZ lower branches pendant. Branchlets slender, firm,
reddish brown, purplish or ‘mahogany’ (variable),
shallowly grooved, glabrous, minutely pubescent (in
Ecology
grooves) or densely pubescent on 1st year shoots; leaf
This rare species has been found on Mt. Fanjing scars circular. Vegetative buds ovoid-oblong, (4–)5–
(Wuling Shan) at 2100–2300 m a.s.l. The climate on 8 × 4–5 mm, slightly resinous; bud scales triangular
this mountain is cool and moist, with only a short ovate, yellowish brown, persistent for several years.
summer season at this altitude. The species occurs Leaves spirally arranged, crowded in several over-
there in a mixed forest with, among other species, lapping ranks of unequal length, the lower leaves
Tsuga chinensis, Acer flabellatum, Rhododendron pectinate, on coning shoots assurgent, (1–)1.5–3(–
hypoglaucum, Enkiartnus chinensis and Prunus ser- 4.5) cm long, 2–3.5 mm wide, twisted at base, ligu-
rulata. Only a few Abies trees are found scattered in late-linear or linear, flat or margins slightly revolute;
the forest. shining green above, whitish or glaucous green be-
low; apex entire, emarginate or bifid, obtuse or acute
on cone bearing shoots. Stomata in two bands di-
Conservation
vided by a midrib on the lower side of leaves. Pollen
Only known from a single locality with a small cones crowded, axillary near shoot apex, small, yel-
population (but the number of mature trees is un- low with red microsporophylls. Seed cones lateral,
known) in forest dominated by other tree species. erect; peduncles very short; cones ovoid-oblong or
This fir should be considered Endangered under cri- oval-cylindrical, with obtuse or umbilicate apex, 5–9
terion D (fewer than 250 mature individuals). Mt. cm long, 3–4 cm wide; bluish purple when imma-
Fanjing has a forest reserve which includes this spe- ture, maturing to purplish or reddish brown. Seed
cies, but we do not know if all the individuals are scales cuneate-flabellate, length × width at mid-cone
within the reserve. 0.8–1.2 × 1.5–2 cm; surface smooth, puberulent; up-
IUCN: EN (D) per margin rounded, entire; base pedicellate. Bracts
obovate-cuneate, 1–1.5 cm long, slightly exserted,
straight or with recurved cusps. Seeds oblong, 5–6
Uses
× 3–3.5 mm, light brown or blackish; seed wings
Although potentially a timber tree, this species does cuneate, oblique, 6–7 × 4–5 mm, light purplish to
not appear to be exploited at present. nearly black.

Distribution
Abies fargesii Franch., J. Bot. (Morot) 13 (8): 256.
1899. Western China: S Gansu, W Hubei, Shaanxi, NW
Sichuan.
TDWG codes: 36 CHC-HU CHC-SC CHN-GS CHN-
Etymology
SA
This species commemorates the French abbé
P. G. Farges, who collected it in 1893.
 gins, apices of leaves on non-coning shoots emar-
Ecology
ginate to bifid. Seeds dark brown to blackish; wings
Abies fargesii occurs in the high montane to subalpine nearly black.
zones of N Central China, at elevations between
2000 m and 4000 m a.s.l. (A. fargesii var. faxoniana
Distribution
at elevations between 2600 m and 4000 m, with an
optimum between 3400 m and 3800 m according to China: S Gansu, W Hubei, Shaanxi, NW Sichuan.
Wang, 1961). Soils are mostly grey brown mountain TDWG codes: 36 CHC-HU CHC-SC CHN-GS CHN-SA
podzols. The climate is cold and moist. At its lowest
elevation broad-leaved trees (e.g. Fagus engleriana,
Conservation
82 Davidia involucrata) are important, but A. fargesii
mostly forms either pure forests or mixed conifer- IUCN: LC
ous forests with among other species Picea purpurea,
P. asperata, P. neoveitchii, P. brachytyla, Larix potani- Abies fargesii Franch. var. faxoniana (Rehd. &
nii, Abies chensiensis, A. recurvata, Tsuga chinensis E. H. Wilson) T. S. Liu, Monogr. Gen. Abies: 151.
and Taxus chinensis. Some broad-leaved trees are 1971. Abies faxoniana Rehd. & E. H. Wilson, in
usually present: Betula spp., Populus spp., and many Sargent, Pl. Wilson. 2: 42. 1914; Abies delavayi
shrubs: Cotoneaster, Ribes, Spiraea, Rhododendron Franch. var. faxoniana (Rehd. & E. H. Wilson)
and Berberis are among the common genera (except A. B. Jacks., in Chittenden, Cult. Conif.: 246. 1932.
in dense Picea-Abies forest). Type: China: Sichuan, Min River, Songpan, Ne of
Songpan, E. H. Wilson 4060 (holotype A).
Uses
Description
Being the most widespread of firs in the high moun-
tains of western China, this species has been subject New shoots densely pubescent with ferruginous
to extensive exploitation for its timber. The wood, short hairs; buds 4–5 cm long, 4 mm wide; leaves
if of high grade, is used in construction (mainly in- with flat margins and emarginate apex. Seeds light
door flooring, framing and joinery), otherwise it is brown, wings pale, tinged purple.
applied in the paper pulp industry. Most botanical
collectors active in western China in the first dec-
Distribution
ades of the 20th century encountered it or its varie-
ties and the species has been introduced to Europe China: W Sichuan.
and the USA from these collections. Most trees still TDWG codes: 36 CHC-SC
in cultivation date from these introductions and are
generally confined to botanical gardens and arbo-
Conservation
reta.
IUCN: LC
3 varieties are recognized:
Abies fargesii Franch. var. sutchuenensis Franch.,
Abies fargesii Franch. var. fargesii. Type: China: J. Bot. (Morot) 13 (8): 256. 1899. Abies sutchuenensis
Chongqing Municipality, Chengkou Xian, Daba (Franch.) Rehd. & E. H. Wilson, in Sargent, Pl.
Shan, [“Tsjen-keou-tin” (district)], P. G. Farges Wilson. 2: 48. 1914. Type: China: Sichuan, Jiange
908bis (holotype P). Xian, Chengkou, P. G. Farges s.n. (holotype P).

Abies fargesii Franch. var. hupehensis Silba, Phytolo- Abies kansouensis Bordères & Gaussen, Trav. Lab.
gia 68: 15. 1990. Forest. Toulouse T. 1 (4, 5): 6. 1944.

Description Description
New shoots glabrous or minutely pubescent; buds New shoots glabrous or minutely pubescent; buds
6–8 mm long, 4–5 mm wide; leaves with flat mar- 6–8 mm long, 4–5 mm wide; leaves with slightly
revolute margins, apices of leaves on non-coning two broad bands separated by a midrib below, none
shoots entire to acute. or a few in a central groove above. Pollen cones soli-
tary in leaf axils, pendulous, 2.5–3 cm long, yellow.
Seed cones lateral, erect; peduncles short, with many
Distribution
scales at base; shape ovoid-oblong or conical, with
China: S Gansu, NW Sichuan. obtuse apex, 8–15 cm long, 3–5 cm wide; yellowish
TDWG codes: 36 CHC-SC CHN-GS green or green with yellow bracts when immature,
maturing to greyish green and becoming yellowish
brown when ripe; cone rachis persistent, narrowly
Conservation
conical, brown. Seed scales broad flabellate, length
IUCN: LC × width at mid-cone 2–2.5 × 2.8–3.2 cm; surface 83
smooth or longitudinally striated, glabrous or spar-
ingly pubescent on exposed parts; upper margin
Abies firma Siebold & Zucc., Fl. Japon. 2 (2): 15, thinner than the rest of the scale, incurved, entire or
t. 107. 1842. Type: Japan: [“in Japonia”], P. F. von obscurely denticulate; base short pedicellate. Bracts
Siebold comm. 1842 ex herb. Zuccarini No. 284 oblanceolate, 2 cm long, exserted, especially on low-
(lectotype M). er part of cone, straight. Seeds obovate-cuneate, 6–8
mm long, light brown; seed wings broad cuneate,
10–15 mm long, yellowish brown.
Etymology
The species epithet (Latin: firmus = firm, stable) re-
Distribution
fers to the rigidity of the leaves.
Japan: Honshu, Kyushu, Shikoku, Yakushima.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Vernacular names
Momi fir; momi, momi-no-ki (Japanese)
Ecology
Abies firma occurs on hills and in mountains of
Description
southern and central Japan, at elevations between
Trees to 50 m tall, d.b.h. to 2 m; trunk monopodi- 50 m and 1900 m a.s.l. (commonly 300 m and 1000
al, massive, straight, columnar, terete; crown open, m). The soils are various mountain soils of volcanic
broad pyramidal, domed or flat topped in old trees. origin or alluvial, and mesic. The climate is moist or
Bark of young trees smooth, with horizontal resin wet, cool in the north of its range and warm tem-
blisters, pinkish grey, in old trees thick, corky, rough, perate in the south, with annual precipitation above
fissured at base. Branches of first order long, spread- 1000 mm. This species is a constituent of mixed
ing horizontally or, especially in young trees, as- forests (rarely in pure stands on dry sites) with e.g.
cending; branches of second order thick, spreading Fagus crenata, F. japonica, Castanea crenata, Carpi-
horizontally or ascending. Branchlets firm, yellow- nus laxifolia, Quercus spp., Tsuga sieboldii, Pinus
ish grey, buff grey brown or light brown, grooved, parviflora, P. densiflora, Pseudotsuga japonica, Abies
usually quite glabrous or with fine pubescence in homolepis, Cryptomeria japonica, Sciadopitys verti-
grooves of 1st year shoots; leaf scars circular or angu- cillata, Chamaecyparis obtusa, Torreya nucifera and
lar. Vegetative buds ovoid to conical, maximal 10 × 5 Picea jezoensis subsp. hondoensis.
mm, not or only slightly resinous; bud scales broad
conical, pale (reddish) brown with green margins,
Conservation
persisting several years. Leaves spirally arranged,
pectinate in 2–3(–4) ranks of different length, on IUCN: LC
coning shoots spreading, the upper leaves assurgent,
(1–)1.5–3.5(–5) cm long, 2–4 mm wide, narrowed and
Uses
curved or twisted at base, linear, flattened; lustrous
(light) green above, whitish green below; apex ob- Momi is the most common and widespread fir in
tuse or emarginate (bifid in young trees). Stomata in southern Japan and there regarded as an ­important
 timber tree. Its wood is light, soft and straight- cm long, yellowish, with purple microsporophylls.
grained and easily worked, but requires careful sea- Seed cones lateral, erect on short peduncles, thick,
soning to prevent warping. It is used for carpentry barrel shaped or cylindrical, with obtuse or retuse
making indoor framing, flooring, joinery, crates, apex, 6–10(–14) cm long, 4–5(–6) cm wide, pur-
boxes etc., but the greatest quantities of its timber plish blue with blue bracts when immature, ripen-
are converted to paper pulp. In plantation forestry it ing to purplish brown or dark brown; cone rachis
is only common in Japan, where old growth stands persistent, massive, cylindro-conical or fusiform,
of this large fir have mostly been logged. Elsewhere, purplish brown. Seed scales cuneate-obovate, length
it is only used as an ornamental tree or planted in × width at mid-cone 2 × 1.8 cm; surface smooth or
collections in arboreta and botanic gardens, requir- somewhat rough, puberulent; upper margin entire
84 ing a climate with mild winters and abundant rain- or slightly erose; base pedicellate. Bracts oblong or
fall. broadly cuneate-spathulate, 2–3 cm long, with long
cusps (–10 mm), much exserted, included or only
the cusps exserted, with straight or recurved cusps.
Abies forrestii Coltm.-Rog., Gard. Chron., ser. 3, 65: Seeds obovate, ca. 8 mm long, lustrous brown; seed
150. 1919. wings cuneate-dolabriform, 10 × 8 mm, light brown.

Etymology Taxonomic notes

This species was named after the English plant hunt- Abies chengii Rushforth was described from cul-
er George Forrest (1873–1932). tivated trees in the British Isles, derived from seed
collected by George Forrest “somewhere in Yunnan”
and was accepted as a species in my book Pinaceae
Vernacular names
(Farjon, 1990). It is more probably another variety
Forrest fir; chuandian lengshan (Chinese) (or a mere form not deserving a botanical name)
of A. forrestii. A hybrid origin with this species and
A. chensiensis subsp. salouenensis has also been sug-
Description
gested, but without supporting evidence. Abies for-
Trees to 40 m tall, d.b.h. to 1–1.5 m; trunk monopo- restii is evidently a variable species.
dial, straight, columnar, terete; crown broad conical,
flat topped in old trees. Bark in young trees smooth,
Distribution
brown grey, in old trees dark brown, fissured at base.
Branches of first order long, spreading horizontally, China: SW Sichuan, NW Yunnan, SE Xizang [Ti-
ascending near the top; branches of second order bet].
assurgent. Branchlets thick, purplish, reddish or TDWG codes: 36 CHC-SC CHC-YN CHT
orange brown, often turning grey, smooth or finely
grooved, glabrous or (ferruginous) pubescent; leaf
Ecology
scars circular or oval. Vegetative buds globular to
ovoid, 4–10 × 3–7 mm, very resinous; bud scales This species (and its varieties) occurs in the high
ovate, red brown (covered with white resin), persist- mountains of SW China at elevations between 2400
ing several years. Leaves spirally arranged, dense, m and 4300 m a.s.l. (commonly 3000–4000 m), on
covering shoot in several ranks, parting in the mid- grey-brown mountain podzols. The climate is cold
dle, assurgent on coning shoots, (1.5–)2–3(–4) cm and wet, annual precipitation ranges from 1000
long, 2–2.5 mm wide, curved or twisted at base, mm to 2000 mm. The species forms forests in pure
linear or ligulate-linear, flattened; margins flat or stands near the tree limit, or is mixed with Picea li-
slightly revolute; lustrous dark green or bluish green kiangensis, Larix potaninii, Tsuga dumosa and some
above, two white (sometimes tending to greenish broad-leaved trees, e.g. Betula albo-sinensis, Acer
white) bands below; apex emarginate, or acuminate spp. and Sorbus spp. at lower elevations. An erica-
on coning shoots. Stomata in two bands separated ceous lower shrub layer with Rhododendron spp. is
by a green midrib below. Pollen cones lateral, 3–4.5 often prominent.
 Abies forrestii Coltm.-Rog. var. ferreana (Bordères
Uses
& Gaussen) Farjon & Silba, Phytologia 68: 17.
Forrest fir and and its several varieties occur at high 1990. Abies ferreana Bordères & Gaussen, Trav.
altitudes, often up to the tree line and consequent- Lab. Forest. Toulouse T. 1 (4, 15): 8. 1947. Type:
ly only yield timber suitable for saw mill process- China: Yunnan, Chungtien, Yua-Tse, T. T. Yü 12326
ing from larger trees at its lowest altitudinal range. (holotype TLF).
Exploitation has (at least officially) ceased with
Chinese forest conservation law now prohibiting Abies chayuensis W. C. Cheng & L. K. Fu, Acta Phy-
logging in old growth forest in the western provin­ totax. Sin. 13 (4): 83. 1975; Abies forrestii Coltm.-Rog.
ces. Having been collected on numerous occasions var. chayuensis (W. C. Cheng & L. K. Fu) Silba, Phy-
by the famous European plant hunters of the early tologia 68: 16. 1990. 85
20th century it was introduced to Europe and the Abies ferreana Bordères & Gaussen var. longibracte-
United States where it is still quite common in ar- ata L. K. Fu & Nan Li, Novon 7 (3): 261. 1997.
boreta and private large gardens. Most trees labeled
A. delavayi ­actually belong to this species (A. del-
Description
avayi has narrow leaves with revolute margins and
dark violet-blue or purplish black seed cones). Re- Shoots pubescent. Resin canals in leaves medial.
newed collecting in recent decades has brought in Seed cones 6–8 cm long.
some new stock on a limited scale. Abies forrestii is
one of the most attractive species of fir in horticul-
Distribution
ture and deserves to be made much more widely
available. Perhaps China could do some alternative China: NW Yunnan, SE Xizang [Tibet].
business now that they have ceased logging? TDWG codes: 36 CHC-YN CHT

4 varieties are recognized:

Conservation
Abies forrestii Coltm.-Rog. var. forrestii. Abies IUCN: LC
delavayi Franch. var. forrestii (Coltm.-Rog.)
A. B. Jacks., in Chittenden, Cult. Conif.: 245. 1932. Abies forrestii Coltm.-Rog. var. georgei (Orr) Far-
Type: China: Yunnan, Lijiang Shan, eastern slopes jon, Pinaceae (Regnum Veg. 121): 59. 1990. Abies
at 27° 25´ N, G. Forrest 6744 (holotype E). georgei Orr, Notes Roy. Bot. Gard. Edinburgh 18: 1,
t. 236. 1933; Abies delavayi Franch. var. georgei (Orr)
Abies chengii Rushforth, Notes Roy. Bot. Gard. Ed- Melville, Bull. Misc. Inf. R. B. G. Kew 1958: 533.
inburgh 41: 333. 1983; Abies forrestii Coltm.-Rog. var. 1959. Type: China: Yunnan, Jinsha-Mekong Divide,
chengii (Rushforth) Silba, Phytologia 68: 17. 1990. G. Forrest 22547 (holotype E).

Description Description
Shoots glabrous. Resin canals in leaves marginal, Shoots densely pubescent with ferruginous, short
small. Bracts slightly exserted, or only the cusps ex- hairs; leaves 1.5–3 cm long; margins slightly revolute.
serted, or sometimes entirely included. Bracts of seed cones exserted, broad and gradual-
ly tapering to a short cusp, with erose-denticulate,
light brown margins.
Distribution
China: SW Sichuan, NW Yunnan, SE Xizang [Ti-
Taxonomic notes
bet].
TDWG codes: 36 CHC-SC CHC-YN CHT In Flora of China 4: 49 (1999) this taxon is treated as
a species, as originally described by M. Y. Orr.
Conservation
IUCN: LC
 Distribution Description
China: SW Sichuan, NW Yunnan, SE Xizang [Ti- Trees to 15 m tall, d.b.h. to 0.5 m; trunk monopodial,
bet]. straight, columnar, terete; crown narrowly conical,
TDWG codes: 36 CHC-SC CHC-YN CHT often open and irregular on exposed sites, broader
in solitary trees. Bark of young trees smooth, brown,
with numerous horizontal resin blisters, becoming
Conservation
rough, scaly and grey in old trees. Branches of first
IUCN: VU (A1cd; A2cd) order long, spreading horizontally or ascending near
the top; branches of second order slender, spreading
86 Abies forrestii Coltm.-Rog. var. smithii Viguié & horizontally or assurgent. Branchlets slender, stiff,
Gaussen, Trav. Lab. Forest. Toulouse T. 1 (2, 1): 177. pale yellowish brown, becoming darker brown in
1929. Abies delavayi Franch. var. smithii (Viguié & the second year, slightly grooved, densely and rather
Gaussen) T. S. Liu, Monogr. Gen. Abies: 143. 1971; persistently pubescent with short, reddish hairs; leaf
Abies georgei Orr var. smithii (Viguié & Gaussen) scars circular. Vegetative buds broadly ovoid, 4 × 3
W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): mm, very resinous; bud scales dark reddish brown,
63. 1975. Type: China: Yunnan, Lijiang Shan, E but covered with yellowish resin, persistent several
slope, J. F. Rock 10673 (holotype A). years. Leaves spirally arranged, the lower ones pec-
tinate, the upper ones spreading upward and for-
ward, incurved and assurgent especially on coning
Description
shoots, 1–2(–2.3) cm long, 2–2.2 mm wide, twisted
Shoots pubescent. Bracts of seed cones exserting, or curved at base, linear or falcate linear, widest near
broad and abruptly narrowing to a long cusp. the obtuse or acute (rarely emarginate) apex, dark
green above, two whitish bands below. Stomata in
two bands separated by a midrib below, a few in the
Distribution
central groove above. Pollen cones crowded on the
China: NW Yunnan. lower side of branches, pendulous, 1 cm long, yellow,
TDWG codes: 36 CHC-YN with red microsporophylls. Seed cones lateral, erect,
often crowded, on short peduncles, oblong-conical,
with narrowed, obtuse apex, 4–8 cm long, 2.5–4 cm
Conservation
wide, dark purple with yellowish bracts when im-
IUCN: LC mature, ripening to purplish brown with pale brown
bracts; cone rachis persistent, narrowly conical,
dark brown. Seed scales broadly flabellate to reni-
Abies fraseri (Pursh) Poir. in Lamarck, Encycl. form, length × width at mid-cone 0.8–1.2 × 1–1.5 cm;
Suppl. 5: 35. 1817. Pinus fraseri Pursh, Fl. Amer. Sept. surface smooth, puberulent on exposed parts; upper
2: 639. 1814; Abies balsamea (L.) Mill. subsp. fraseri margin entire and incurved; base long pedicellate.
(Pursh) E. Murray, Kalmia 12: 18. 1982. Type: not Bracts oblong or obovate-oblong, with obcordate
designated. apices and short cusps, 1.5–2 cm long, much exsert-
ed and reflexed, covering most of the seed scales.
Seeds obovate-cuneate, 5–6 mm long, dark purplish
Etymology
black; seed wings obliquely cuneate or dolabriform,
This species was named after John Fraser (1750–1811) 5–6 × 5 mm, purplish brown.
who collected plants in eastern North America,
among which was this fir.
Distribution
USA: W North Carolina, E Tennessee, SW Virginia
Vernacular names
(Appalachian Mts.).
Fraser fir, Fraser’s fir TDWG codes: 78 NCA TEN VRG
 Abies grandis (Douglas ex D. Don) Lindl., Penny
Ecology
Cyclop. 1: 30. 1833. Pinus grandis Douglas ex
On the highest slopes and summits of the Appala- D. Don, in Lambert, Descr. Pinus, ed. 8°, 2: p. s.n.
chian Mountains, between 1200 m and 2038 m a.s.l., inter 144 et 145 Type: not designated.
usually best developed on N-facing slopes. The soils
are commonly podzolized and moderately acid. Abies excelsior Franco, Bol. Soc. Brot., ser. 2, 23: 162.
The climate is humid, with cool summers and cold 1949.
winters with heavy snowfall, annual precipitation Abies grandis (Douglas ex D. Don) Lindl. var. ida-
varies between 850 mm and 2000 mm. Fraser fir hoensis Silba, Phytologia 68: 19. 1990.
occurs in scattered populations, sometimes pure at
the highest elevations, but more often mixed with 87
Etymology
Picea rubens and Betula papyrifera above 1500 m, at
lower elevations also with Tsuga caroliniana, Betula The species epithet is a reference to the great stature
alleghaniensis, Sorbus americana, Acer saccharum this species can attain in its natural habitat.
and Fraxinus caroliniana. Ericaceae and various
herbs are common in the understorey, often thick
Vernacular names
moss carpets (Hylocomium splendens) cover the for-
est floor. Grand fir, Lowland fir, Giant fir

Conservation Description
The disjunct populations of this fir, restricted to the Trees to 80–100 m tall, d.b.h. to 2–3 m; trunk mono­
mountain tops and their north-facing slopes of the podial, straight, columnar, terete; bole often free
southern Appalachians, are susceptible to destruc- of branches to considerable height; crown nar-
tion by windfall and fire. However, by far the most rowly conical. Bark of young trees thin, smooth
damaging agent is an insect, the Balsam woolly and grey brown, with many resin blisters, braking
adelgid (Adelges piceae), discovered in 1957 in Abies up into many small plates in old trees. Branches of
fraseri on Mt. Mitchell. This alien pest has spread first order long, spreading horizontally, the lower
quickly to all populations causing massive dieback ones curved downward; branches of second order
through impairment of translocation flow in the spreading, or ascending near the end of the main
cambium. Millions of trees had died by the 1980’s branches. Branchlets slender, firm, olive green to
and only one substantial population (Mt. Rogers, reddish brown, with narrow, straight ridges between
Virginia) remained largely unaffected. Methods the leaves, minutely pubescent, but glabrous by the
to control this introduced insect are still being re- 3rd year; leaf scars small, circular. Vegetative buds
searched but none have been fully effective; some globose, 1.5–2 × 1.5 mm, slightly resinous; bud scales
small scale protection can be provided by chemical triangular, obtuse, red-brown, persisting 2–3 years.
insecticides. Leaves spirally arranged, pectinately spreading in
IUCN: VU (D2) a more or less horizontal plane, the upper ranks
shorter than the lower, on coning shoots assurgent,
(2–)3–5(–6) cm long, 2–3 mm wide, strongly twist-
Uses
ed at base, narrowly linear, flattened with slightly
The remaining stands of Fraser fir have very limited revolute margins, dark glossy green above, greenish
commercial value as timber trees. The most impor- white below; apex emarginate or obtuse on coning
tant use is growing this species for Christmas trees; shoots. Stomata only on the abaxial (lower) surface
it is considered the best conifer available in the USA in two bands separated by a midrib. Pollen cones lat-
for this purpose. It has a natural ‘Christmas tree eral on the lower sides of branches, axillary, 1.2–1.8
shape’ and retains its fragrant, dark green leaves cm long, yellowish green. Seed cones lateral, erect;
well for this indoor use. It is also widely used as an peduncles very short (cones nearly sessile); shape
ornamental tree for gardens with several cultivars oblong-­cylindrical, with obtuse apex, (5–)7–12 cm
named. At least in the UK it does not usually have a long, 3–4 cm wide, light green or purple tinged when
very long life as a garden tree. immature, ripening to dull grey-brown, ­usually very
 resinous; cone rachis persistent, narrowly conical, because they tend to grow up very symmetrically and
brown. Seed scales broad flabellate, length × width have lustrous green foliage. Grand fir is commonly
at mid-cone 2–2.5 × 2.5–3 cm; surface smooth, in grown as an amenity tree in large gardens and city
immature cones pubescent; upper margin entire, parks and, as another David Doulas introduction, it
incurved; base pedicellate. Bracts short, rectangular, was planted in nearly all landscape gardens laid out
with very small cusps, 1–1.5 cm long, entirely includ- in the 19th century in Europe, where some trees have
ed. Seeds cuneate, 8 × 5 mm, pale brown; seed wings now attained impressive sizes. In horticulture it is
cuneate-dolabriform, 10–15 mm long, pale brown much in use and a substantial number of cultivars
with a purple tinge. have been selected for garden planting.

88
Distribution
Abies guatemalensis Rehd., J. Arnold Arbor. 20:
SW Canada, NW USA: south to N California. 285. 1939.
TDWG codes: 71 BRC 73 IDA MNT ORE WAS 76 CAL
Etymology
Ecology
The species epithet refers to Guatemala, from where
Grand fir has its optimum in lowland coastal areas it was first described.
of the Pacific Northwest, but occurs also in the Cas-
cade Range and the northern Rocky Mountains, to
Vernacular names
the west of the Continental Divide. It grows from
near sea level to ca. 1800 m a.s.l., on a variety of Guatemalan fir
soils derived from granitic or basaltic rock, best de-
velopment is on alluvial soils with a relatively high
Description
ground water table. In the Pacific Northwest the
climate is moist maritime to wet, with annual pre- Trees to 35–40 m tall, d.b.h. to 1–1.5 m; trunk mo-
cipitation from as low as 500 mm to 2500 mm, in nopodial, straight, columnar, terete; crown broad
the upland interior the winters are snowy and cold, conical or dome shaped and open in old trees. Bark
the precipitation ranges from 500 mm to 1250 mm. in young trees smooth, grey brown, becoming scaly
It grows pure in some areas in Idaho, but is usually in old trees. Branches of first order long, spread-
mixed with Pseudotsuga menziesii, Abies amabilis, ing horizontally, the lower ones curved downward;
Picea sitchensis, Calocedrus decurrens, Thuja plicata, branches of second order spreading. Branchlets
Tsuga heterophylla or Larix occidentalis (in the in- slender, greenish to reddish brown or purplish red,
terior). Broad-leaved associated trees are e.g. Acer turning dark purple, finally dark grey, ridged and
macrophyllum, Alnus rubra (along streams), and grooved, with minute pubescence or glabrous; leaf
Fraxinus latifolia, while the shrub layer is formed scars ovate or circular. Vegetative buds globose to
by Acer circinatum in coastal areas and Amelanchier ovoid, 4–5 × 3–3.5 mm, resinous; bud scales broadly
­alnifolia and Rosa spp. in the interior. triangular ovate, with obtuse apex, reddish brown,
persisting 2–3 years. Leaves spirally arranged, in two
lateral sets, pectinate to subdistichous, at right an-
Conservation
gles to shoot or obliquely forward, on coning shoots
IUCN: LC the upper (shorter) leaves assurgent, (1.2–)1.5–5(–7)
cm long, 1.2–2 mm wide, twisted at base, narrowly
linear, flattened, with emarginate or sometimes ob-
Uses
tuse apex, shining green, with two white bands be-
Rapid growth and great size make this species an im- low. Stomata absent above or a few near apex, in two
portant timber tree. The wood is soft and white and bands separated by a midrib below. Pollen cones
an excellent source of pulpwood. For construction lateral, 2–2.5(–3.5?) cm long, yellowish. Seed cones
timber it is considered less desirable due to its rela- lateral, erect, subsessile, oblong-cylindrical, with ob-
tive weakness and limited durability. In the Pacific tuse or truncate apex, 8–12(–16) cm long, 4–5.5 cm
Northwest young trees are valued as Christmas trees wide, purple when immature, becoming dark brown
when ripe; cone rachis persistent, narrowly conical, is therefore only of limited importance as a timber
blackish brown. Seed scales transversely oblong to tree. Like A. hickelii, it is locally worked in sawmills
reniform, length × width at mid-cone 2–2.5 × 3 cm; for domestic use in carpentry, plywood and veneer.
surface smooth, puberulent; upper margin erose- In Guatemala, there is trade in the foliage branches
denticulate or entire, slightly revolute; base pedicel- for semi-natural Christmas trees and the species
late. Bracts cuneate-obovate, with short triangular is also grown in plantations for that purpose. Its
cusps, 1.2–1.7 cm long, included or slightly exserted. southern distribution has made it an unlikely tree to
Seeds cuneate-obovoid, 8–9 mm long, light brown; be tried in horticulture in Europe and North Ameri-
seed wings broad, obliquely obovate, 10–15 × 10–15 ca, but provenances from the highest altitudes could
mm, light brown. be more or less hardy, certainly in the SW USA and
southern Europe, where it is, however, still rarely 89
seen even in botanical collections. A few trees grow
Distribution
in botanic gardens in England and in California.
El Salvador; Guatemala; Honduras; S Mexico along
the Sierra Madre del Sur to Jalisco and Zacatecas. 2 varieties are recognized:
TDWG codes: 79 MXE-TA MXS-GR MXS-JA MXS-
MI MXS-OA MXT-CI 80 ELS GUA HON Abies guatemalensis Rehd. var. guatemalensis.
Type: Guatemala: Huehuetenango, Las Cumbres
del Aire, J. H. Faull 13104 (holotype A).
Ecology
Abies guatemalensis is the southernmost species of Abies tacanensis Lundell, Amer. Midl. Naturalist 23:
the genus: it reaches 14° 49´ N in the Guatemalan 175. 1940; Abies guatemalensis Rehd. var. tacanensis
highlands. It grows from 1800 m to 3700 m a.s.l. (Lundell) Martínez, Pinaceas Méxic., ed. 3: 129. 1963.
on the Pacific side of the Sierra Madre del Sur, in Abies zapotekensis Debreczy, Rácz & Ramírez, Phy-
Guatemala usually between 3300 m and 4100 m, on tologia 78 (3): 9. 1995.
well drained mountain soils of volcanic origin. The Abies guatemalensis Rehd. var. longibracteata Debre-
climate is cool, moist oceanic, with most precipita- czy & Rácz, Phytologia 78 (3): 11. 1995.
tion as rain in the winter, or as fog the year round
in Guatemala. Var. guatamalensis occurs between
Description
3500 m and 3800 m a.s.l. in the mountains of Chia-
pas, at a greater distance from the ocean than the New shoots at first purple, turning dark grey. Leaves
other varieties. Guatemalan fir is usually associated (1.2–)1.5–4(–5.5) cm long. Pollen cones 2–2.5 cm
with several highland pines, e.g. Pinus ayacahuite, long when shedding pollen. Seed cones 8–12 cm
P. montezumae, P. hartwegii, P. pseudostrobus and long, 4–5.5 cm wide; bracts shorter or as long as the
P. devoniana, also with Abies religiosa in W Guate- seed scales, minutely cuspidate.
mala and with Cupressus lusitanica in Chiapas. At
lower elevations Juniperus spp., Quercus spp. and
Distribution
Arbutus spp. become more important.
El Salvador; W Guatemala; Honduras; Mexico from
Chiapas to Nayarit and Tamaulipas.
Conservation
TDWG codes: 79 MXE-TA MXS-CL MXS-GR MXS-
Listed on CITES Appendix I. JA MXS-MI MXS-NA MXS-OA MXT-CI 80 ELS GUA
HON
Uses
Conservation
Guatemalan fir occurs in scattered stands on high
mountains, often in quite inaccessible places, and IUCN: VU (A1d)
 Abies guatemalensis Rehd. var. jaliscana Martínez, dish brown, ridged and grooved between the leaves,
Anales Inst. Biol. Univ. Nac. México 19: 73. 1948. glabrous or with minute pubescence in the grooves;
Type: Mexico: Jalisco, Talpa de Allende, Cuale, Las leaf scars small, ovate or circular. Vegetative buds
Mesas, M. Martínez 28000 (holotype MEXU). ovoid or globular, 5 × 4 mm, very resinous; bud
scales triangular ovate, brown, but covered in yel-
Abies religiosa (Kunth) Schltdl. & Cham. var. emar- lowish resin, persisting several years. Leaves spirally
ginata Loock ex Martínez, Anales Inst. Biol. Univ. arranged, pectinate, in two lateral sets at about right
Nac. México 19: 60. 1948. angles to shoot, of about equal length or the upper
Abies flinckii Rushforth, Notes Roy. Bot. Gard. Edin- ranks shorter, on coning shoots the upper leaves
burgh 46: 101. 1989. assurgent, (1.2–)1.8–3.5 cm long, 1–1.8 mm wide,
90 twisted or curved at base, shining light green above,
glaucous below; apex emarginate or sometimes ob-
Description
tuse. Stomata in two bands separated by a midrib
New shoots initially greenish, turning light reddish below, none or a few near apex above. Pollen cones
brown. Leaves 3.5–7 cm long. Seed cones 9–14(–16) lateral, short, yellow with red microsporophylls.
cm long, (3–)4–5 cm wide; bracts or bract cusps ex- Seed cones lateral, erect, short pedunculate, oblong-
serted. Pollen cones possibly longer than in var. gua- cylindrical, with obtuse apex, 6–8(–12) cm long, 2.5–
temalensis, but insufficiently known. 3.5(–5) cm wide, purple when immature, ripening to
dark brown; cone rachis persistent, cylindric-coni-
cal, blackish brown. Seed scales cuneate-flabellate,
Distribution
length × width at mid-cone 1.3–1.5 × 1.8–2 cm; sur-
Mexico: Jalisco. face smooth, hirsute-puberulent; upper margin
TDWG codes: 79 MXS-JA rounded, entire; base pedicellate. Bracts oblong-lan-
ceolate, 2 cm long, exserted, straight. Seeds obovate-
cuneate, 6–7 mm long, light brown; seed wings
Conservation
broad cuneate, 10 × 8 mm, light brown.
IUCN: VU (A1d)
Taxonomic notes
Abies hickelii Flous & Gaussen, Trav. Lab. Forest. In a recent paper Strandby et al. (2009) presented
Toulouse T. 1 (1, 17): 1. 1932. a morphometric study of the genus Abies in Mexi-
co and Central America. They proposed to reduce
A. hickelii to a subspecies of A. religiosa on the basis
Etymology
of their results. This paper came too late for these
This species was named after the French botanist results to be thoroughly considered (or tested) and
and dendrologist R. Hickel. therefore this new taxonomy is merely noted here.

Vernacular names Distribution

Hickel’s fir; oyamel, pinabete (Mexican Spanish) Mexico: Chiapas, Guerrero, Oaxaca.
TDWG codes: 79 MXS-GR MXS-OA MXT-CI
Description
Ecology
Trees to 30 m tall, d.b.h. to 1–1.3 m; trunk monopo-
dial, straight, columnar, terete; bole often bare of Both varieties of this species occur in high moun-
branches to a considerable height; crown narrowly tains of sub-tropical S Mexico, at elevations between
conical, or wider and more open in old trees. Bark of 2500 m and 3000 m a.s.l. The soils are of volcanic
young trees smooth, grey, of old trees thick, break- origin. The climate is cool, moist oceanic, with rain
ing into quadrangular plates. Branches of first order mostly in the winter. There are some pure stands
spreading horizontally; of second order spreading at the highest elevations, but this species is usually
or ascending. Branchlets slender, purplish or red- mixed with highland pines, e.g. Pinus montezumae,
P. pseudostrobus, and P. ayacahuite, and also with
Conservation
Cupressus lusitanica and Quercus spp. Shrubs are
e.g. Vaccinium spp., Andromeda spp., Ribes spp. and IUCN: VU (A1d)
Fuchsia spp.

Abies hidalgensis Debreczy, Rácz & Guízar, Phy­

Uses
tologia 78 (3): 4. 1995. Type: Mexico: Hidalgo,
Hickel’s fir is a rare species and its two varieties oc- Metepec, 4–5 km N of the village, Z. Debreczy et al.
cur in more or less disjunct, limited stands. Exploi- DAPC 40323 (holotype BP).
tation for timber is minor and its use is local, mainly
worked in sawmills for domestic purposes. In cul- 91
Etymology
tivation it is extremely rare and limited to some
dendrological collections in countries with mild cli- This species was named after the Mexican State
mate, e.g. southern France. of Hidalgo; the name also recalls Miquel Hidalgo
y Costilla (1762–1811), after whom the State was
2 varieties are recognized: named.

Abies hickelii Flous & Gaussen var. hickelii. Type:

Vernacular names
Mexico: Oaxaca, Sierra de San Felipe (?), 1900,
C. Conzatti s.n. (holotype LY). No common name has been given to this species.

Description Description
Seed cones 6–8(–9) cm long, 2.5–3.5 cm wide. Trees monopodial; trunk columnar, straight (max.
height and diam. not given); crown columnar-coni-
cal. Bark smooth, light grey in young trees, on large
Distribution
trunks breaking into irregular, large plates, thick;
Mexico: Chiapas, Guerrero, Oaxaca. inner bark red. Branches spreading and ascend-
TDWG codes: 79 MXS-GR MXS-OA MXT-CI ing, higher orders descending in lower part of the
tree. Foliage branchlets robust, smooth or ridged
and grooved on leading shoots, new shoots densely
Conservation
pubescent with short, yellowish brown hairs. Veg-
IUCN: VU (A1d) etative buds small, partly hidden by short, incurv-
ing leaves, slightly resinous; bud scales triangular.
Abies hickelii Flous & Gaussen var. oaxacana Leaves pectinately arranged, distichous, shortest on
(Martínez) Farjon & Silba, Phytologia 68: 20. 1990. the upperside of branchlets, (1–)3–5(–6) cm long,
Abies oaxacana Martínez, Anales Inst. Biol. Univ. 1.5 mm wide, linear, curving slightly downward or
Nac. México 19: 39. 1948. Type: Mexico: Oaxaca, forward, shortly bifid or emarginate at apex, lus-
M. Martínez 29000 (holotype MEXU). trous dark green or dull greyish green, with white
stomatal bands on the abaxial side; stomata in few
Abies hickelii Flous & Gaussen var. macrocarpa Mar- to several lines on the upperside, in two white bands
tínez, Anales Inst. Biol. Univ. Nac. México 13 (2): below separated by a green midrib; resin canals 2–5,
621. 1942. medial, in leaves on coning branches up to 7. Pol-
len cones lateral, crowded, 10–12 mm long, yellow,
Seed cones 9–12 cm long, ca. 5 cm wide. with puberulent microsporophylls. Seed cones lat-
eral, erect, short pedunculate, cylindrical, 6.5–8 cm
long, 3.5–4 cm wide, pruinose green at maturity,
Distribution
with obtusely rounded or slightly umbilicate apex;
Mexico: Guerrero, Oaxaca. cone rachis narrowly conical. Seed scales flabellate
TDWG codes: 79 MXS-GR MXS-OA to wide cyathiform, green to greyish green, densely
velutinous with very short, white, appressed hairs.
 Bracts short, 5–8 mm long, spathulate, with serrate Abies holophylla Maxim., Bull. Acad. Imp. Sci.
margins, without a protruding cusp, entirely in- Saint-Pétersbourg 10: 487. 1866. Type: China:
cluded. Seeds narrowly triangular with resin blisters [“Manchuria”], 8 Sep 1869, C. J. Maximowicz s.n.
(size not given); seed wings yellowish grey (not fully (holotype LE).
developed at time of description).
Abies holophylla Maxim. var. aspericorticea Y. Y. Sun,
Bull. Bot. Res. (China) 25 (3): 264. 2005.
Taxonomic notes
The type collection, Debreczy et al. DAPC 40323
Etymology
(holotype BP, isotypes A, E, K, MEXU) was gathered
92 4.5 km N of the village of Metepec. The number of The species epithet means ‘entire leaf ’ and refers to
resin ducts in the leaves, given as 5–7 in the proto- the undivided leaf apex.
logue, is highly unusual in the genus, but A. hickelii
has 4–12. Abies hidalgensis could be nothing more
Vernacular names
than a variety of A. hickelii with included bracts
in the seed cones. It is only known from the type Needle fir, Manchurian fir; shansong (Chinese)
specimens deposited in the herbaria cited above.
More material is needed to establish its taxonomic
Description
position, the species is here given the benefit of the
doubt. Trees to 30–50 m tall, d.b.h. to 1–1.5 m; trunk mo-
nopodial, straight, columnar, terete; crown shape
broad pyramidal, young trees with narrowing top.
Distribution
Bark of young trees smooth, grey to buff orange, in old
Mexico: Hidalgo (canyon near village of Metepec). trees shallowly fissured, scaly and brown. Branches
TDWG codes: 79 MXE-HI of first order long, spreading horizontally; branches
of second order spreading horizontally or assurgent.
Branchlets stout, firm, yellowish to grey-brown or
Ecology
buff orange, with ridges and grooves between the
This species occurs in a steep canyon in the Sierra leaves, glabrous, or with only minute pubescence
Madre Oriental at an altitude of ca. 2000–2300 m in grooves of 1st year shoots; leaf scars oval circular,
a.s.l. on the moist, eastern slopes in cloud forest. It light coloured. Vegetative buds ovoid-conical, 4–8 ×
is associated with Cupressus lusitanica, Pinus patula, 3–5 mm, more or less resinous; bud scales triangular
P. pseudostrobus and P. teocote; among broad-leaved with blunt apices, prominently keeled, light brown
trees Quercus laurina, Ternstroemia pringlei and Ce- or reddish brown, persisting several years. Leaves
strum sp. dominate, with Alnus firmifolia, Buddleia spirally arranged, radial and assurgent on leading
cordata, Ptelea trifoliata and Sambucus mexicanum and coning shoots, subdistichous and/or more or
as frequent understorey trees and shrubs. less pectinate in shaded shoots, 2–4.5 cm long, 2–2.5
mm wide, twisted or curved at base, linear, taper-
ing towards an acute or mucronate apex, flattened,
Conservation
lustrous light green above, pale whitish green with
This species was described and named as new in a green midrib and margins below. Stomata in two
1995 from a single location in Hidalgo, Mexico. Its bands divided by a midrib below, none or a few near
distribution beyond the type locality remains un- apex above. Pollen cones lateral, clustered, 1–1.5 cm
known and no subsequent collections have been as- long, yellow with reddish microsporophylls. Seed
signed to this new species. cones lateral, erect; peduncles 0.5–1 cm, scaly; shape
IUCN: DD oblong-cylindrical, with obtuse or truncate apex,
6–14 cm long, 3–4.5 cm wide, pale green when im-
mature, sometimes with a purple tinge, maturing
Uses
to yellowish green and ripening to light yellow-
No uses are known, or likely, of this species. It has ish brown; cone rachis persistent, narrowly coni-
not been taken into cultivation. cal, brown. Seed scales ­flabellate, length × width
at mid-cone 2 × 3 cm; surface smooth, puberulent and botanic gardens of the northern hemisphere’s
on exposed parts, often very resinous; upper mar- cooler regions.
gin entire, slightly incurved; base pedicellate. Bracts
oblong-spathulate, with small cusps, 0.8–1 cm long,
included. Seeds cuneate, ca. 6 × 4 mm, light brown; Abies homolepis Siebold & Zucc., Fl. Japon. 2 (2):
seed wings cuneate, with a truncate, oblique end, ca. 17, t. 108. 1842. Fig. 8
13 × 10 mm, light brown.
Etymology
Distribution
The species epithet (Greek/Latin: homo = equal, the
NE Asia: from mountains N of Vladivostok to South same; lepis = scale) refers to the similar length of 93
Korea, also in Heilongjiang Prov. and Jilin Prov., seed scales and bracts.
China.
TDWG codes: 31 PRM 36 CHM-HJ CHM-JL 38 KOR-
Vernacular names
NK KOR-SK
Nikko fir; dake-momi, nikko-momi (Japanese)
Ecology
Description
Abies holophylla occurs in NE China in low to medi-
um high mountains, in SE Siberia and N Korea also Trees to 35–40 m tall, d.b.h. to 1–1.5 m; trunk mo-
in hills and on lowland plains. Its elevational range is nopodial, straight, columnar, terete; crown broad py-
from 10 m to 1200 m a.s.l. in the north, to 500–1500 ramidal, in old trees often flat topped. Bark of young
m a.s.l. in the south of its range. It grows on lithosols trees smooth, with papery flakes, grey with pinkish
of granitic origin. The climate is cold, with wet sum- hues, of old trees rough and broken into plates, dark
mers and arid winters with long periods of snow. At grey-brown. Branches of first order long, spreading
higher elevations or in the NE of its range it forms horizontally; branches of second order spreading
pure stands, or more commonly mixed coniferous horizontally or assurgent. Branchlets stout, firm,
forests with Pinus koraiensis, especially in the coast- lustrous light yellowish brown, ridged and deeply
al mountains near the Sea of Japan. In other areas grooved, glabrous; leaf scars circular or ovate. Veg-
it is a constituent of the northern mixed coniferous etative buds ovoid-conical, 3–5 × 2–4 mm, resin-
deciduous forests, with Abies nephrolepis, Picea obo- ous; bud scales triangular, keeled, reddish brown,
vata, Larix gmelinii (var. olgensis) and broad-leaved persisting several years. Leaves spirally arranged,
trees, such as Populus spp., Quercus mongolica, Frax- more or less distichous, but pectinate on shaded
inus mandshurica, Ulmus spp., and Betula ermanii. shoots, assurgent to recurved on coning shoots, but
usually parted above, (1–)1.5–3(–3.5) cm long, 2–3.5
mm wide, slightly twisted or curved at base, linear
Conservation
or ligulate-linear, flattened, deeply grooved and lus-
IUCN: LC trous green above, two whitish bands below; apex
obtuse or emarginate. Stomata in two broad bands
separated by a midrib below. Pollen cones lateral,
Uses
crowded, pendulous, yellow. Seed cones lateral,
Manchurian fir is a valuable timber tree in its native erect, often several together; peduncles 0.5 cm; shape
area and is exploited mainly in managed forests in oblong-cylindrical, with obtuse or umbilicate apex,
much of the region. Most of its wood today is used 7–10(–14) cm long, 2.5–3.5(–5) cm wide, violet blue
for plywood and veneer as it is evenly grained, light or greenish when immature, violet-brown or brown
and easily worked. Minor uses are for soundboards when ripe; cone rachis persistent, narrowly coni-
in musical instruments, boxes and sometimes for cal, blackish brown. Seed scales broadly flabellate to
joinery. In Europe it was introduced around 1905, flabellate, length × width at mid-cone 1.5–2 × 2.5–3
but, although perfectly hardy and ornamental, re- cm; surface smooth, puberulent, often very resin-
mains an uncommon tree mainly seen in arboreta ous; upper margin entire, incurved; base pedicel-
late. Bracts rectangular-oblong, 1 cm long, included.
 Seeds cuneate, 6 × 4 mm, light brown; seed wings
Distribution
cuneate-dolabriform, 10 × 8 mm, yellowish brown.
Japan: central Honshu, Kyushu, Shikoku.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Distribution
Japan: central Honshu, Kyushu, Shikoku.
Conservation
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
IUCN: LC
Ecology
Abies homolepis Siebold & Zucc. var. umbellata
94 Abies homolepis is a species of high mountains in (Mayr) E. H. Wilson, Conif. Tax. Japan: 58. 1916;
the central parts of the Japanese islands Honshu, Abies umbellata Mayr, Monogr. Abiet. Japan. Reich.:
Kyushu and Shikoku. In the south it occurs from 34, t. 1, f. 2. 1890; Abies brachyphylla Maxim. var.
1100 m to 1800 m a.s.l., in Honshu between 700 umbellata (Mayr) Dallim. & A. B. Jacks., Handb.
m and 2000 m. The soils are mesic, derived from Conif.: 88. 1923. Type: not designated.
volcanic rock and usually well drained. The climate
is cool and humid. Near the tree limit it forms ei-
Description
ther pure stands, or mixtures with A. veitchii and/
or Larix kaempferi, but at lower elevations it occurs Seed cones 10–14 cm long, 4–5 cm wide, greenish;
in the mixed coniferous deciduous forests, with e.g. apex more or less umbilicate.
Fagus crenata, Quercus crispula, Betula grossa, Tsuga
diversifolia, Thuja standishii, Pinus densiflora, Picea
Distribution
jezoensis subsp. hondoensis; A. firma replaces A. ho-
molepis below 1100 m. Japan: central Honshu, Kyushu, Shikoku.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Uses
Conservation
Nikko fir is not an important timber tree as its range
is limited and it occurs at high altitudes. It is fairly IUCN: LC
widely planted as an ornamental tree in Japan as well
as in Europe, where it appears to be one of the least
demanding species in the genus. A few cultivars are Abies kawakamii (Hayata) T. Itô, Encycl. Japon.
known in Japan as well as in Europe (independently 2: 167. 1909. Abies mariesii Mast. var. kawakamii
derived), mostly being dwarfed forms suitable for Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25 (19): 223.
Japanese gardens or rockeries. 1908. Type: Taiwan: Nantou, Chia-i Pref., Yu-Shan,
[“Mt. Morrison”], T. Kawakami & U. Mori 2369
2 varieties are recognized: (lectotype TI). Fig. 9

Abies homolepis Siebold & Zucc. var. homolepis.

Etymology
Type: Japan: [“in Japonia”], P. F. von Siebold comm.
1842 ex herb. Zuccarini No. 290 (lectotype M). The species epithet commemorates the Japanese
plant collector T. Kawakami.
Abies brachyphylla Maxim., Bull. Acad. Imp. Sci.
Saint-Pétersbourg 10: 488. 1866; Abies firma Siebold
Vernacular names
& Zucc. var. brachyphylla (Maxim.) Bertrand, Bull.
Soc. Bot. France 18: 380. 1871. Taiwan fir; Taiwan lengshan (Chinese)

Description Description
Seed cones 7–10 cm long, 2.5–3.5 cm wide, violet- Trees to 16–20 m tall, d.b.h. to 0.5–1 m; trunk mo-
blue; apex obtuse. nopodial, straight, terete, in some old trees bent, but
otherwise erect; crown broad pyramidal, on moun- of the wettest mountain ranges in the world. There
tain tops irregular and open. Bark of young trees are some pure forests on N and NE slopes at these
smooth, soon flaking, light grey or yellowish grey, in high elevations (3200 m to 3600 m a.s.l.), or the spe-
old trees becoming rough and scaly, greyish brown. cies occurs mixed with scattered Pinus armandii var.
Branches of first order heavy and long, spread- mastersiana, Tsuga chinensis var. chinensis, ­Picea
ing horizontally or curved; branches of second or- morrisonicola, and with Juniperus squamata var.
der spreading or assurgent. Branchlets stout, firm, morrisonicola at the upper limit of Abies. At lower
pale yellowish brown, with prominent ridges and elevations the forest becomes progressively more
grooves, light brown pubescence in grooves, on cone mixed with broad-leaved trees, e.g. Acer insulare,
bearing shoots pubescence red brown; leaf scars cir- Trochodendron aralioides, Quercus semecarpifolia
cular. Vegetative buds conical-ovoid, 4–5 mm long, subsp. glabra, Ilex bioritsensis, and Eurya spp. Other 95
very resinous; bud scales triangular, keeled, purple- conifers in this belt are Tsuga chinensis var. chinensis,
brown, persisting several years. Leaves spirally ar- which becomes more abundant than Abies kawaka-
ranged, spreading radially, pectinate below shoot, mii between 2400 m and 3000 m a.s.l., Pseudotsuga
shorter and assurgent above shoot, covering it en- sinensis, and Chamaecyparis obtusa var. formosana,
tirely, on coning shoots assurgent, (0.8–)1–2.2(–2.5) which is more abundant below 2400 m (Liu, 1971).
cm long, 1.5–2 mm wide, slightly twisted or curved
at base, linear-falcate, flattened, lustrous green
Conservation
above, whitish green below; apex emarginate or ob-
tuse, acuminate on cone bearing shoots. Stomata Logging of this species, which occurred mainly dur-
in two narrow bands separated by a midrib below. ing the period of Japanese occupation (1895–1945),
Pollen cones lateral, crowded, 1 cm long, yellowish. has ceased almost completely and substantial popu-
Seed cones lateral, erect, short pedunculate, oblong- lations now occur within national parks and other
cylindrical, with obtuse or umbilicate apex, (3–)5– reserves. Its limited distribution and occurrence in
7(–9) cm long, 2.5–4 cm wide, dark purple when a mozaic with subalpine bamboo grassland makes
immature, dark purple-brown when ripe; cone ra- it vulnerable to fires that could be caused by much
chis persistent, cylindric-conical or narrowly fusi- increased tourism.
form, dark purplish brown. Seed scales flabellate, IUCN: NT
length × width at mid-cone 1.5–1.7 × 2–2.3 cm; sur-
face smooth, puberulent; upper margin erose; base
Uses
pedicellate. Bracts obcordate-rectangular, with very
small cusps, 1–1.3 cm long, included. Seeds cuneate- The timber of this species was formerly exported
oblong, 6–8 mm long, blackish brown; seed wings to Japan, where it was used for general carpentry. It
cuneate (oblique), 6–10 mm long, light brown with is little used for this purpose today in Taiwan. This
a black tinge. species was introduced to England in 1930 and is oc-
casionally found in arboreta in Europe and North
America, but remains uncommon in cultivation.
Distribution
Taiwan (central mountains).
TDWG codes: 38 TAI Abies koreana E. H. Wilson, J. Arnold Arbor. 1:
188. 1920. Type: South Korea: Cheju-do, Halla-san,
[“Hallai-san Quelpart Island”], E. H. Wilson 9486
Ecology
(holotype A). Fig. 10
A high mountain species, occurring between 2400
m and 3800 m a.s.l. in the central high mountains Abies koreana E. H. Wilson f. nigrocarpa Hatus.,
of Taiwan, on grey brown podzolized soils and also Bull. Kyushu Univ. Forest. 5: 40. 1934.
on mountain yellow earth, both acid and usually
rocky. The climate is temperate, super humid: above
Etymology
humid subtropical foothills the annual precipitation
exceeds 4000 mm, with maxima up to 10,000 mm, The species name denotes its origin, Korea where
making the Taiwanese central high mountains one Ernest Wilson discovered it.
 cuneate-dolabriform, 4–6 mm long, light brown,
Vernacular names
tinged with purple.
Korean fir
Distribution
Description
South Korea (Cheju Island, Chiri-san, Halla-san,
Trees to 15–18 m tall, d.b.h. to 0.5–0.8 m; trunk mo- Kaya-san, Kongo-san, Mudung-san, Tokyu-san).
nopodial, straight, columnar, terete; crown broad TDWG codes: 38 KOR-SK
pyramidal. Bark of young trees smooth, thin, with
resin blisters, light grey with a purplish tinge, in old
Ecology
96 trees becoming thick and rough, breaking into ir-
regular plates near base, greyish black. Branches On the medium high mountains of South Korea it
of first order densely set to low above the ground, occurs only at the higher elevations and summits,
spreading horizontally, the lower curved downward; on shallow mountain soils poor in humus content.
branches of second order spreading, ascending near Its elevational range is between 1000 m and 1900 m
the top. Branchlets slender, firm, yellowish grey or a.s.l. The climate is cool temperate, with a summer
grey-green, shallowly grooved between the leaves, monsoon bringing the annual precipitation above
sparsely pubescent in the grooves, soon glabrous; 1600 mm. Abies koreana grows in pure stands or
leaf scars circular. Vegetative buds subglobose, 4–5 mixed with Betula ermanii on Cheju Island; on the
× 3–4 mm, very resinous; bud scales obtuse, mem- mainland it is also mixed with Picea jezoensis, Pi-
branous, brown, persisting several years. Leaves spi- nus koraiensis, Taxus cuspidata, Quercus mongolica
rally arranged, radially spreading, the upper leaves var. mandshurica, Cornus controversa, Acer spp., and
shorter and curved upward or recurved, especially several genera of low shrubs, e.g. Juniperus, Deutzia,
so on coning shoots, (0.8–)1–2(–2.2) cm long, 2–2.5 Ribes and Rhododendron. The forest is usually open
mm wide, slightly twisted or curved at base, linear- and essentially dominated by conifers, of which
spathulate (often widest near the apex), flattened, A. koreana is often a minor component.
with slightly revolute margins, lustrous dark green
above, two white bands and green midrib and mar-
Conservation
gins below; apex emarginate, obtuse or sometimes
acute. Stomata none or a few near the apex above, IUCN: NT
in two broad bands separated by a midrib below.
Pollen cones lateral, clustered around shoot, 1 cm
Uses
long, yellowish with scarlet microsporophylls. Seed
cones lateral, erect, often crowded together, numer- Korean fir is a small to medium sized tree that grows
ous, even on the lower branches; peduncles 0.4–0.6 slowly and together with its rarity these qualities
cm; shape cylindrical, with obtuse, umbilicate or make it unsuitable as a timber tree. It is, however,
papilliform apex, 4–7 cm long, 2.5–3 cm wide, pur- perhaps the most widely used species in the genus
ple or violet blue with greenish bracts when im- for gardens. Its small stature, compact growth and
mature, maturing to purplish brown, with light the appearance of decorative seed cones even in very
brown bracts, ripening to dull brown or purplish small trees have made it a target of growers for the
brown; cone rachis persistent, narrowly conical, selection of several cultivars. Wilson’s original intro-
dark brown. Seed scales reniform, or wing-shaped duction came from Cheju Island (Quelpart Island)
(Liu, 1971, p. 45), length × width at mid-cone 1–1.2 × where trees remain small; later introductions from
1.2–1.6 cm; surface smooth, pubescent; upper mar- mainland Korea have produced better growing trees
gin erose, incurved; base pedicellate. Bracts broadly and, after nearly a century some have reached good
spathulate-obcordate, 1.2–1.5 cm long, exserted, size. The colours of bracts and seed scales of cones
(strongly) reflexed. Seeds cuneate, 4–6 mm long, vary much and are one of the attractions of this
light or dark brown, with a purple tinge; seed scales ­conifer.
Abies lasiocarpa (Hook.) Nutt., N. Amer. Sylva 3: e­ ntire, curved inward; base pedicellate. Bracts ob-
138. 1849. long, with small cusp, 0.6–1 cm long, included.
Seeds cuneate-oblong, 5–6 mm long, brown; seed
wings cuneate-oblong, with oblique end, 10–18 mm
Etymology
long, light brown tinged with purple.
The species epithet describes the strongly pubescent
(hairy) ‘fruits’ (i.e. seed cones).
Distribution
W North America: from Yukon to New Mexico, Ari-
Vernacular names
zona and N California.
Subalpine fir, Alpine fir TDWG codes: 70 ASK NWT-MK YUK 71 ABT BRC 97
73 COL IDA MNT ORE WAS WYO 76 ARI CAL NEV
UTA 77 NWM
Description
Trees to 30–35 m tall, d.b.h. to 1–1.2 m; trunk mo-
Ecology
nopodial, straight, columnar, terete; crown narrowly
conical, forming a graceful spire. Bark of young trees This is a species of the subalpine zone in the high
smooth, thin, with horizontal resin blisters, white or mountains of W North America, occurring from
grey, in old trees becoming thicker, creamy white 5 m to 1500 m a.s.l. in the north of its range and
or grey, fissured and brown or grey towards base. between 600 m and 3500 m in the Cascade Range
Branches of first order very dense, down to near and Rocky Mountains. It grows on a variety of high
the ground, short, spreading horizontally, pendant mountain lithosols, wet or dry. The climate is eve-
below, the ends curved upward; branches of second rywhere cold, but humid in the NW and dry in the
order spreading. Branchlets slender, pliable, very S of its range, precipitation varies between 500 mm
strong, pale brown, soon grey, with prominent ridg- and 3000 mm annually. It forms usually very open
es and grooves, pubescent with short brown hairs; stands with solitary or clustered trees, often mixed
leaf scars circular. Vegetative buds globose or sub- with Tsuga mertensiana in the NW and with Picea
globose, 3.5–6 × 3–5 mm, very resinous; bud scales engelmannii in most of the Rocky Mountains. Other
obtuse, keeled, brown, persisting several years. conifers are mainly Pinus spp., and also Abies spp.
Leaves spirally arranged, the lower longest and pec- in the Pacific Northwest. Alpine meadows typically
tinate, the other leaves radially spreading, the upper occur between the clumps of conifers.
leaves curved forward and assurgent, especially so
on coning shoots, 2.5–4.5 cm long, 1.5–2 mm wide,
Uses
slightly twisted at base, narrowly linear, curved or
bent near base, flattened, longitudinal groove on the Subalpine fir has little or no importance as a tim-
upper surface shallow, glaucous green, with 2 whit- ber tree even though its wood properties are gener-
ish bands below; apex obtuse to acute or weakly ally similar to other species of fir. To a large extent
emarginate. Stomata in several rows in the adaxial this is of course due to the rich heritage of conifers
groove, in two bands separated by a strong midrib western North America enjoys above many other
below. Pollen cones lateral, usually crowded, pen- parts of the world; there is no need to use them as
dulous, 1.5–2 cm long, yellow, with purplish blue timber trees. Subalpine fir also tends to grow slowly,
microsporophylls. Seed cones lateral, erect, shortly remains relatively small and grows in inaccessible
pedunculate or sessile, oblong-cylindrical, with ob- places. It may be used locally for construction tim-
tuse apex, 7–10 cm long, 2.5–3.5 cm wide, purple, or ber, doors, window frames, boxes and other such
purplish green with greyish pubescence when im- products, but the wood tends to be knotty due to the
mature, becoming purplish brown or brown with retention of branches. Although it naturally grows
yellowish pubescence when ripe; cone rachis per- into the perfect Christmas tree shape, it is rarely
sistent, narrowly conical, dark brown. Seed scales used as such, because it grows slowly and cutting it
cuneate-trapeziform, length × width at mid-cone from its habitat is environmentally destructive. It is
1.5–2.5 × 1.3–2.3 cm; surface smooth, strongly pu- also little used in horticulture (except perhaps cul-
bescent with yellowish brown hairs; upper ­margin tivars derived from var. arizonica) for taking it into
 the lowlands of temperate regions usually exposes it
Distribution
to damage from ‘late’ frosts.
USA: Arizona, Colorado, New Mexico.
2 varieties are recognized: TDWG codes: 73 COL 76 ARI 77 NWM

Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa.

Ecology
Pinus lasiocarpa Hook., Fl. Bor. Amer. 2 (10): 163.
1838; Abies balsamea (L.) Mill. subsp. lasiocarpa Var. arizonica is most commonly mixed in forests
(Hook.) Boivin, Naturaliste Canad. 86 (10): 223. with Picea engelmannii, Pinus aristata and P. flexi-
1959. Type: USA: Washington, Columbia River, lis, or it occurs in pure stands, at elevations between
98 D. Douglas s.n. (holotype K). 2400 m and 3650 m a.s.l.

Abies subalpina Engelm. var. fallax Engelm., Trans.

Conservation
St. Louis Acad. Sci. 3: 597. 1878; Abies lasiocarpa
(Hook.) Nutt. var. fallax (Engelm.) Franco, Abetos: IUCN: LC
15. 1950.

Abies magnifica A. Murray bis, Proc. Roy. Hort.

Description
Soc. London 1863 (3): 318. 1863. Fig. 11
Bark on large trunks relatively thin, grey-brown, fis-
sured towards base.
Etymology
The species epithet means ‘magnificent’ or ‘out-
Distribution
standing’ or any equivalent adjective.
W North America: from Yukon to New Mexico and
N California.
Vernacular names
TDWG codes: 70 ASK NWT-MK YUK 71 ABT BRC 73
COL IDA MNT ORE WAS WYO 76 CAL NEV 77 NWM Red fir, California red fir

Conservation Description
IUCN: LC Trees to 60–70 m tall, d.b.h. to 2–3 m; trunk mo-
nopodial, massive, straight, columnar, terete; crown
Abies lasiocarpa (Hook.) Nutt. var. arizonica narrowly conical, in old trees open. Bark of young
(Merriam) Lemmon, Bull. Sierra Club 2: 167. trees smooth, with resin blisters, grey or light pur-
1898. Abies arizonica Merriam, Proc. Biol. Soc. plish grey, in old trees thick, rough, deeply fissured
Washington 10: 116. 1896; Abies balsamea (L.) Mill. below, brown. Branches of first order relatively
subsp. lasiocarpa (Hook.) Boivin var. arizonica short, spreading, the lower bent downward or pen-
(Merriam) Boivin, Naturaliste Canad. 86 (10): dant; branches of second order spreading horizon-
223. 1959; Abies lasiocarpa (Hook.) Nutt. subsp. tally, pendant at last. Branchlets slender, strong but
arizonica (Merriam) E. Murray, Kalmia 12: 18. 1982; pliable, light brown or greenish, with ridges and
Abies bifolia A. Murray bis var. arizonica (Merriam) grooves between leaves, for about two years densely
O’Kane & K. D. Heil, Harvard Pap. Bot. 7 (2): 324. pubescent with reddish brown hairs; leaf scars circu-
2003. Type: not designated. lar. Vegetative buds ovoid globose or more conical,
small, hidden by terminal short leaves, not resinous;
bud scales triangular, keeled, brown puberulent, per-
Description
sisting 2–3 years. Leaves spirally arranged, spreading
Bark on large trunks thick, creamy white, increas- laterally, but curved inward above shoot, on coning
ingly fissured towards base. shoots assurgent and strongly curved, concealing
shoot, 2–3.5 cm long, 1.3–1.6 mm wide, (abruptly)
curved at base, narrowly linear, curved, only slightly
flattened, no groove above, glaucous grey green on tion per ha even in natural, unmanaged stands. It is
both sides; apex obtuse or acute. Stomata in two sep- therefore increasingly valuable as a timber tree used
arate rows of lines above, in two bands separated by a for general construction and plywood. This species
midrib below. Pollen cones lateral, crowded, pendu- is also valued as a Christmas tree, both grown in
lous, 1.5–2 cm long, with scarlet microsporophylls. natural stands and in plantations. It is relatively rare
Seed cones lateral, erect, short pedunculate or ses- in amenity plantings with few cultivars known; most
sile, broad conical or barrel shaped, with truncate existing planted trees date from the heydays of land-
or umbilicate apex, (10–)13–20 cm long, (5–)7–10 scape conifer plantings in the 19th century.
cm wide, purplish green when immature, matur-
ing to yellowish brown or greenish brown, ripen- 2 varieties are recognized:
ing to light brown; cone rachis persistent, narrowly 99
conical, dark brown. Seed scales cuneate-flabellate, Abies magnifica A. Murray bis var. magnifica.
length × width at mid-cone (2.5–)3–4 × (2.5–)3–4 Type: USA: California, Sierra Nevada, W. Lobb 441
cm; surface smooth, densely pubescent with yellow- (holotype K).
ish brown hairs; upper margin entire and incurved;
base long pedicellate. Bracts oblong-spathulate, with
Description
a short cusp, 2–3.5 cm long, included or exserted and
reflexed. Seeds cuneate-oblong, 13–15 × 5–6 mm, Seed cones 14–20 cm long, 7–10 cm wide; bracts in-
dark brown; seed wings cuneate, 15–20 mm long, cluded.
lustrous roseate brown.
Distribution
Distribution
USA: California, W Nevada, SW Oregon.
USA: California, W Nevada, SW Oregon. TDWG codes: 73 ORE 76 CAL NEV
TDWG codes: 73 ORE 76 CAL NEV
Conservation
Ecology
IUCN: LC
Abies magnifica occurs in the Canadian Life Zone of
high mountains, between 1400 m and 2700 m a.s.l. Abies magnifica A. Murray bis var. shastensis
(to 3000 m in the south of its range); commonly on Lemmon, [Cone-bearers Calif.] Calif. State Board
soils of granitic (Sierra Nevada) or basaltic (Cascade Forest. Bienn. Rep. 3: 145. 1890. Abies shastensis
Range) origin, which have been altered by glaciation (Lemmon) Lemmon, Gard. & Forest 10: 184. 1897.
and are usually slightly acid. The climate is charac- Type: not designated.
terized by short, warm and dry summers and long,
cold winters with much snow. Annual precipitation
Description
varies between 750 mm and 1500 mm (80 % as snow).
This species forms pure stands in some places, but Seed cones 10–13 cm long, 5–8 cm wide; bracts ex-
more often it is a constituent of the mixed conifer- serted and reflexed.
ous forest type with e.g. Pinus spp., Abies concolor,
A. procera, Pseudotsuga menziesii, Calo­cedrus decur-
Taxonomic notes
rens, Juniperus occidentalis, and at higher elevations
Abies lasiocarpa and Tsuga mertensiana subsp. gran- Opinions vary among authors: Liu (1971) and Silba
dicona. Common shrubs are e.g. Ceanothus cordu- (1986) have treated this variety as a natural hybrid
latus, Chrysolepis sempervirens and Arctostaphylos between Abies magnifica and A. procera, Rushforth
nevadensis. (1987) kept it as a variety of A. magnifica and Lit-
tle (1979) treated it as a synonym of A. magnifica. It
seems to occur within the northern range of A. mag-
Uses
nifica and the extreme southern range of A. procera
California red fir grows to large dimensions with ex- and ‘transitional’ cone types are found in a broad
tremely straight boles and has a high wood produc- zone with its southern limit around Lassen Peak
 in N California (R. J. Laacke in Burns & ­Honkala, resinous; bud scales brown, ciliate, persistent for 2–3
1990). Exserted bracts are a character state of years. Leaves spirally arranged, the lower leaves pec-
A. procera and A. magnifica var. magnifica has only tinate, the upper ones curved inward and forward,
slightly smaller cones than that species, unlike the shorter, covering shoots, on cone bearing branches
substantially smaller cones of var. shastensis. These almost erect, (0.6–)1–2(–2.5) cm long, 1.6–2.5 mm
character states are probably not constant within wide, widest near apex, strongly twisted and nar-
A. magnifica as there is a population of the latter in rowed at base, linear, the upper leaves more ligulate
the southern Sierra Nevada, well beyond the tran- and curved, flattened, grooved above, with slightly
sition zone, that shows exserted bracts. Artificial revolute margins and emarginate apex (obtuse on
crossing experiments indicate the possiblity of in- coning shoots), lustrous dark green above, 2 white
100 trogression via pollen from A. procera into A. mag- bands below separated by green midrib. Stomata in
nifica. two bands separated by a midrib on lower surface
only. Pollen cones lateral, axillary, pendulous, 1.5–2
cm long, yellowish. Seed cones lateral, short pedun-
Distribution
culate or sessile, ovoid-oblong, with obtuse apex,
USA: from Lassen Peak in California to Crater Lake 4–9 cm long, 2–4.5 cm long, violet-blue, ripening
in Oregon. to dark blackish purple (brown inside); cone rachis
TDWG codes: 73 ORE 76 CAL persistent, narrowly conical, blackish brown. Seed
scales cyathiform-flabellate, length × width at mid-
cone 1.5–2.2 × 2–2.5 cm; surface smooth, puberulent
Conservation
on exposed parts; upper margin entire, incurved;
IUCN: LC base pedicellate. Bracts obovate or obcordate, 1–1.5
cm long, included. Seeds conical-ovoid, (4–)5–7
mm long, light brown; seed wings cuneate, 10–12 × 8
Abies mariesii Mast., Gard. Chron., ser. 2, 12: 788. mm, light brown, with a purplish tinge.
1879. Type: Japan: Honshu, Awomori Pref., Nikko-
san, C. Maries 73 (holotype K). Pl. 2
Distribution
Japan: Honshu.
Etymology
TDWG codes: 38 JAP-HN
This species was named after C. Maries, who col-
lected it on Mt. Nikko (Mt. Hakkoda) in 1878.
Ecology
A species of the high mountain sides and ridges in
Vernacular names
the upper montane and subalpine zones, occurring
Maries’ fir; O-shirabiso (Japanese) commonly between 1000 m and 2800 m a.s.l. (as low
as 750 m in N Honshu). The soils are mostly derived
from volcanic rock, usually podzolic and slightly
Description
acid or neutral, well drained, and moderately moist
Trees to 25–30 m tall, d.b.h. to 0.8–1 m; trunk mo- (mesic). The climate is cold, with abundant winter
nopodial, straight, columnar, terete; crown broad snow and cool, moist summers, the annual precipi-
pyramidal, flat topped in old trees. Bark of young tation exceeds 2000 mm in the mountains nearest to
trees smooth, pale grey, nearly white, scaly and the Sea of Japan. Frequent typhoons are a destruc-
rough near base and dark grey in old trees. Branch- tive force reducing the maximum age of trees. Abies
es of first order long, spreading wide, soon curved mariesii forms sometimes pure forests near the tree
down to pendant; branches of second order spread- line, but is more common in mixed (coniferous) for-
ing horizontally, dense. Branchlets firm, slender, ests with e.g. Abies veitchii, Tsuga diversifolia, Picea
light brown, ridged and grooved between the leaves, jezoensis var. hondoensis and/or undergrowth of Pi-
densely pubescent with brown hairs, but soon gla- nus pumila and Juniperus communis var. nipponica,
brous and light grey; leaf scars circular. Vegetative the latter two especially abundant on ridgetops.
buds globose, covered by apical leaves, 2–3 mm long, Common broad-leaved trees are Betula ermanii,
 3
6

101

4 5

Pl ate 2 . Abies mariesii. 1. Habit of tree. 2. Foliage branch. 3. Seed cone. 4, 5. Seed scales with bract (4),
with seeds (5). 6. Rachis of seed cone. 7. Leaves.
 Sorbus commixta, and Acer spp. In many previously Bark of young trees smooth, light grey, becoming
disturbed areas with deep, fine textured soil, e.g. vol- rough and scaly with age, fissured at base. Branches
canic ash, there is a dense cover of small bamboo of first order spreading horizontally; branches of
(Sasa paniculata and S. nipponica), which excludes second order idem. Branchlets stout, firm, yellow-
most other plants (Franklin et al., 1979). ish green, shiny, turning grey, with prominent ridges
between the leaves, glabrous, or minutely pubescent
in the first year; leaf scars circular, with a light cen-
Conservation
tral part. Vegetative buds conical, 8–9 × 4–6 mm,
IUCN: LC slightly resinous; bud scales triangular-ovate, lac-
iniate, protruding, light brown, persisting several
102 years. Leaves spirally arranged, spreading laterally
Uses
in two sets, or the upper leaves covering shoot and
This species of fir has little value as a timber tree be- directed forward, the lower leaves pectinate on
cause it grows at high altitude and mostly in inacces- shaded shoots, leaves on coning shoots assurgent,
sible localities. In horticulture it is rather uncommon (1–)1.5–2(–2.2) cm long, 2–3.5 mm wide, twisted or
despite its attractive dark green foliage leaves and curved at base, linear-ligulate, flattened, longitudi-
contrasting white stomatal bands underneath. It is nally grooved above, bright, lustrous green on the
not at all tolerant of droughts and performs best in adaxial (upper) surface, two greenish white bands
cool, wet conditions but on light, well-drained soils. on the lower surface; apex variable: acute or mucro-
It is mostly restricted to collections in botanic gar- nate, on shaded shoots obtuse but not emarginate.
dens and arboreta. Stomata none or a few near the apex above, in two
bands separated by a midrib below. Pollen cones lat-
eral, crowded, 1.5–2 cm long, greenish yellow with
Abies nebrodensis (Lojac.) Mattei, Boll. Reale Orto purple microsporophylls. Seed cones lateral, erect,
Bot. Palermo 7: 64. 1908. Abies pectinata Gilib. var. short pedunculate, cylindrical, with a conical apex,
nebrodensis Lojac., Fl. Sicula 2 (2): 401. 1904–1907; (7–)8–10(–12) cm long, 3–4 cm wide, yellowish
Abies alba Mill. var. nebrodensis (Lojac.) Svoboda, green when immature, ripening to greenish brown
Trudy Bot. Inst. Akad. Nauk SSSR., ser. 1, Fl. Sist. (light reddish brown inside); cone rachis persistent,
Vyss. Rast. 13: 60. 1964; Abies alba Mill. subsp. narrowly conical, brown. Seed scales cuneate-cyat-
nebrodensis (Lojac.) Nitz., Lustgården 1968: 178. hiform, length × width at mid-cone 2–2.5 × 2.8–3.3
1969. Type: not designated. Fig. 12 cm; surface smooth or slightly wrinkled, puberulent
on exposed parts; upper margin entire or undulate;
Abies pectinata Guss., Fl. Sicula Syn. 2: 614. 1844, non base pedicellate. Bracts linear-spathulate, 2.5–3 cm
Gilib. (1792). long, exserted and recurved. Seeds conical-oblong,
6–8 mm long, reddish brown; seed wings cuneate,
oblique, 10–15 mm long, light brown.
Etymology
The species epithet refers to the Monti Nebrodi, an
Distribution
alternative name for the Madonie Mountains where
this species was discovered. Italy: Sicily (Madonie Mountains: Monte Scalone,
Polizzi Generosa).
TDWG codes: 13 SIC-SI
Vernacular names
Sicilian fir
Ecology
A very local remnant population occurs in high
Description
mountains at around 1500 m a.s.l., on calcareous,
Trees to 10–15 m tall in Sicily at present, but may rocky soil. Summers are warm and dry, winters mild
grow taller, d.b.h. to 0.4–0.6 m; trunk monopodial, and moist, the annual precipitation is 700 to 800
straight, columnar, terete; crown (broad) conical (the mm. No real forest stand is left of this once abun-
only living old tree in the wild had its top broken off). dant species. It is today restricted to a few scattered
trees associated with secondary maquis, in which genes of other species. As the synonymy given above
Quercus petraea and, at its highest (potential) limit, indicates, this taxon has (not without reason) been
Fagus sylvatica are the most common low trees. Juni- interpreted as merely a variety or a subspecies of
perus communis forms dense ground covering car- the European fir Abies alba and there is no doubt
pets and could provide protection for seedlings of that it will hybridize with that species if the two are
A. nebrodensis against dehydration and/or grazing brought (once more?) together.
and browsing animals.

Abies nephrolepis (Trautv. ex Maxim.) Maxim.,

Conservation
Bull. Acad. Imp. Sci. Saint-Pétersbourg 10: 486.
Abies nebrodensis is one of the rarest conifers in the 1866. Abies sibirica Ledeb. var. nephrolepis Trautv. 103
world, with only 29 individual plants remaining in ex Maxim., Mém. Acad. Imp. Sci. Saint-Pétersbourg
the wild in 2006. Most of these are small trees grow- (Sav. Etr.) 9: 206. 1859. Type not designated. Pl. 3
ing on heavily overgrazed scree slopes with clumps
of Fagus sylvatica and Quercus petraea scattered Abies ×sibiriconephrolepis Taken. & J. J. Chien, Acta
among them. Extensive logging since the beginning Phytotax. Sin. 6 (1): 153. 1957.
of the 18th century had brought this species to the
brink of extinction when it was ‘discovered’ nearly
Etymology
a century ago in a nearby village, and more recently
the remaining trees in the valley of Madonna degli The species epithet is a Greek word composition: ne-
Angeli on Mt. Scalone. An extensive ex situ con- phrolepis = with kidney-shaped scales.
servation programme both locally and abroad is in
operation but attempts at re-introduction have to
Vernacular names
date not been very successful due to harsh summer
conditions and a totally depleted soil. New attempts Hinggan fir, Khinghan fir; chou lengshan (Chinese);
funded by the EU and providing initial planting Pikhta amurskaya (Russian)
compost and watering in summer may be more
successful; protection from direct sunlight, erosion
Description
and grazing are also essential. Ultimately the forest
ecosystem has to be restored for the survival of this Trees to 30–35 m tall, d.b.h. to 1–1.2 m; trunk mono­
relict conifer. podial, straight, columnar, terete; crown conical
IUCN: CR (D) or oval, old trees with densely branched flat tops
(“stork’s nests”). Bark of young trees smooth, light
grey to greyish brown, in old trees shallowly fis-
Uses
sured, dark greyish brown. Branches of first order
The cultivation of this species in several countries spreading horizontally, those near the top ascend-
of Europe north of the Alps has been very success- ing, the lowest drooping; branches of second order
ful and many trees are now growing well in arboreta spreading horizontally, dense. Branchlets slender,
and other collections. The next phase may well see firm, yellowish grey brown, turning to grey; surface
it develop into a popular amenity tree. While this ridged and grooved, with minute pubescence in the
development is a positive one, at least safeguarding grooves; leaf scars circular. Vegetative buds ovoid or
a threatened species from total oblivion, it is not to conical, broad, 5 × 4 mm, resinous, especially near
be seen as a substitute for conservation of the natu- apex; bud scales triangular, blunt, brown or pur-
ral population. Ex situ conservation, i.e. safeguard- plish red, appressed, persisting several years. Leaves
ing species outside their habitat, is an effort to breed spirally arranged, radially around shoot, directed
individuals with the aim of re-introducing them forward and assurgent, covering the upper part
or their propagules in the wild once the causes of of shoot, 1–2.5(–3) cm long, 2 mm wide, strongly
decline have been eliminated. Most trees planted twisted at base, linear, flattened, with slightly revo-
outside their natural habitat will remain where they lute margins, dull, light (grey-)green, with whitish
are, and their seed, if viable, is without proper pre- green bands below; apex variable: emarginate, ob-
cautions taken usually not guaranteed free from the tuse, acute or acuminate. Stomata none or a few near
 4

104

Pl ate 3. Abies nephrolepis. 1. Habit of trees. 2. Foliage branch. 3. Branch with seed cones. 4. Detail of
foliage. 5, 6. Leaves. 7. Seed scale with bract. 8. Seeds.
apex above, in two bands divided by a midrib below. tain soils. The climate is cold, with short, cool and
Pollen cones lateral, axillary, crowded at the under- moist summers and long, cold winters. Most of the
side of foliage, yellowish green with purple micro- annual precipitation is snow. It is usually associated
sporophylls. Seed cones lateral, erect, often several with other conifers, e.g. Pinus koraiensis and ­Picea
crowded together, short pedunculate, cylindrical, ­jezoensis; also with Pinus pumila and Juniperus
with obtuse apex, 4.5–7.5 cm long, 2–3.5 cm wide, sabina var. davurica at higher elevations (maritime
usually reddish purple and rarely green when im- provinces of the Russian Far East); in the interior
mature, maturing to purplish brown, becoming dull with Picea obovata, Larix gmelinii, Pinus sibirica or
brown when ripe; cone rachis persistent, narrowly Abies sibirica. Betula spp. and Sorbus amurensis are
conical, dark brown with blackish purple tinge. Seed common associated broad-leaved trees.
scales reniform, length × width at mid-cone 1–1.2 105
× 1.5–1.8 cm; surface smooth, often covered with
Conservation
thick clots of yellowish white resin, puberulent on
exposed parts; upper margin entire or slightly erose, IUCN: LC
incurved; base pedicellate. Bracts spathulate, with
prominent, straight cusps, 1.4–2 cm long, only the
Uses
cusps exserted. Seeds obovate-cuneate, 5 × 3 mm,
blackish brown or shining black; seed wings short, Hinggan fir is an important timber tree in NE China
dolabriform, with rounded edges, 6 × 5 mm, black- and Korea. Its wood is used in carpentry and for ply-
ish brown or black. wood and veneer. The relatively small size of this tree
on marginal sites makes exploitation commercially
unlikely; the better, larger trees come from mixed
Taxonomic notes
conifer forests at middle elevations in the moun-
A natural hybrid between A. sibirica and A. nephro­ tains. In horticulture, it was introduced to Great
lepis has been reported from Heilongjiang Province, Britain in 1908 from the botanic garden in St. Pe-
China: Abies ×sibiriconephrolepis Taken. & Chien. tersburg, Russia. It remains a rarely planted species,
Its leaves are described as being shorter than those which is susceptible to damage by late spring frosts
of A. nephrolepis and its young shoots are greyish in countries with an Atlantic maritime climate.
brown pubescent, while the seed cones are larger.
Other character states are intermediate between the
two parent species. It is reported to be common in Abies nordmanniana (Steven) Spach, Hist. Nat.
river valleys in the Lesser Hinggan Range. The oc- Vég. Phan. 11: 418. 1841. Pl. 4
currence of A. sibirica in this region is not confirmed
by other accounts (e.g. Flora of China 4, 1999) and
Etymology
the hybrid status of the firs in this area is doubtful.
This species was named after the botanist A. von
Nordmann, who introduced it to horticulture in
Distribution
1838.
Russian Far East: from the Zeya River to the Sikhote
Alin Range; NE China: Manchuria, Shaanxi, south to
Vernacular names
Hebei (Wutai Shan); North Korea and South Korea.
TDWG codes: 31 AMU KHA PRM 36 CHM CHN-HB Caucasian fir, Nordmann fir; Pikhta kavkazkaya
CHN-SA 38 KOR-NK KOR-SK (Russian)

Ecology Description
This is a species of low to medium high mountains, Trees to 50 m tall, d.b.h. to 1.5–2 m; trunk mono­
occurring at elevations between 500 m and 700 m podial, straight, columnar, terete; crown broad coni-
a.s.l. in E Siberia at the northern limit of its range, cal or pyramidal, old trees often flat topped. Bark of
between 750 m and 2000 m a.s.l. in NE China. This young trees smooth, grey, in old trees rough, shal-
species grows on a variety of well drained moun- lowly fissured, blackish grey-brown. Branches of first
106

6
7

4 5

Pl ate 4. Abies nordmanniana. 1. Habit of tree. 2. Foliage. 3. Seed cone. 4, 5. Seed scales with bract (4),
with seeds (5). 6. Bract. 7. Leaves.
order spreading, ascending in upper part of crown; tinct subspecies of A. nordmanniana, the earliest
branches of second order spreading horizontally, as- combination made at that rank, A. nordmanniana
surgent near the top. Branchlets slender, firm, light subsp. equi-trojani, is the valid name.
olive brown or brown, ridged and grooved, with
brown pubescence in grooves or glabrous; leaf scars
Distribution
circular, light. Vegetative buds ovoid, with pointed
apex, 6 × 5 mm, not resinous or resinous; bud scales Caucasus Mts.; N & W Turkey.
ovate, acute, keeled and with laciniate edges, red- TDWG codes: 33 NCS TCS 34 TUR
brown, persisting several years. Leaves spirally ar-
ranged, pectinate below, the upper leaves pressed
Ecology
forward above shoot, on coning shoots assurgent, 107
(1.5–)2–3(–3.5) cm long, 1.5–2.5 mm wide, strongly Abies nordmanniana is a fir of the high montane
twisted at base, linear, flattened, sometimes with zone of the mountains around the eastern Black Sea,
slightly revolute margins, grooved above, lustrous at elevations between 900 m and 2100 m a.s.l., on
dark green above, two whitish green bands below; soils derived from igneous and granitic rocks. The
apex emarginate, obtuse on coning shoots. Stoma- climate is characterized by warm summers and cold
ta in two bands separated by a midrib below. Pol- winters, with a high annual precipitation, ranging
len cones lateral, crowded, pendulous, 1–2 cm long, in the Caucasus Mountains between 1000 mm and
yellowish. Seed cones lateral, erect, often crowded, 3000 mm. On the mountains Kaz Dagh and Ulu
short pedunculate, ovoid-cylindrical, with pointed Dagh in NW Turkey the subsp. equi-trojani extends
and often papilliform apex, (10–)12–16(–20) cm to the tree line. This species occurs either in pure
long, 4–5.5(–6) cm wide, greenish when immature, stands or mixed with Picea orientalis, at lower eleva-
ripening to light brown (reddish brown inside); cone tions also with Pinus sylvestris, gradually merging
rachis persistent, narrowly conical, dark brown. into deciduous broad-leaved forest with Fagus ori-
Seed scales flabellate or cyathiform, length × width entalis, Acer trautvetteri, Carpinus caucasica, Ulmus
at mid-cone 1.8–2.5 × 2.7–4 cm; surface smooth, elliptica, Acer pseudoplatanus, Tilia caucasica, Taxus
slightly striated, pubescent on exposed parts; up- baccata, and Rhododendron ponticum.
per margin entire, often repand, slightly incurved;
base pedicellate. Bracts spathulate-obcordate, with
Uses
laciniate edges and a short or long cusp with mid-
rib, 2.7–3.5 cm long, exserted and reflexed. Seeds Caucasian fir is an important timber tree in the W
cuneate, 10–12 × 7–8 mm, fawn brown, shiny; seed Caucasus and NW Turkey, where it provides large
wings cuneate, with rounded edge, 15–18 × 15 mm, sizes of valuable, straight grained and easily work-
light purplish brown or rose, turning fawn brown. able wood for building materials, especially veneer.
It grows fast and to large size even in plantations
and parks provided the climate is moist enough.
Taxonomic notes
It is better appreciated in horticulture than other
It is now generally accepted that the firs of NW Tur- large European firs because of regular shape and
key are conspecific with Abies nordmanniana, which dense foliage. Its popularity as a Christmas tree is
has its main distribution in the Caucasus Moun- largely due to its retention of needles well beyond
tains. The disjunct populations of NW Turkey have the time the young trees are normally kept indoors.
been known as and are sometimes still accepted as A large number of cultivars has been selected, both
two distinct species: A. bornmuelleriana (Ulu-Dagh) with variant foliage colours and different branching
and A. equi-trojani (Kaz-Dagh), but the morpho- habits, including dwarf forms and spreading forms
logical differences between the two are minor and without an upright leader shoot.
the character states are overlapping, not discrete. It
seems better to recognize just one subspecies for the Two subspecies are recognized:
populations of NW Turkey. If recognized as a dis-
 Abies nordmanniana (Steven) Spach subsp.
Distribution
nordmanniana. Pinus nordmanniana Steven, Bull.
Soc. Imp. Naturalistes Moscou 11: 45, t. 2. 1838. NW Turkey: Kaz-Dagh (Ida Mts.), Ulu-Dagh (Mt.
Type: Georgia: Caucasus Mts. (South), [“source Olympus of Bithynia).
of the Kur River”], A. von Nordmann s.n. in herb. TDWG codes: 34 TUR
Steven (holotype H).
Conservation
Description
The disjunct populations in NW Turkey here recog-
Vegetative buds not resinous; young foliage shoots nized as A. nordmanniana subsp. equi-trojani have a
108 (branchlets) with brown pubescence in grooves. limited area of occupancy (AOO) compared to the
Cusps on bracts long, with a distinct midrib. species as a whole. The population on Ulu Dagh,
although reasonably safe at present, could suffer
if tourist development (ski resort, roads and lifts)
Distribution
would expand below the tree line in future. Infesta-
Caucasus Mts, N Turkey (Paphlagonia). tion with Dwarf mistletoes (Arceuthobium oxycedri,
TDWG codes: 33 TCS-AB TCS-AD TCS-GR 34 TUR Loranthaceae) has been observed in the fir forest on
this mountain during a visit in 2004.
IUCN: NT
Conservation
IUCN: LC
Abies numidica de Lannoy ex Carrière, Rev. Hort.
Abies nordmanniana (Steven) Spach subsp. equi- 37: 106. 1866. Abies pinsapo Boiss. var. numidica (de
trojani (Asch. & Sint. ex Boiss.) Coode & Cullen, Lannoy ex Carrière) Salomon, Deutsche Bäume
Notes Roy. Bot. Gard. Edinburgh 26: 167. 1965. Sträuch.: 26. 1884; Abies pinsapo Boiss. subsp.
Abies pectinata Gilib. var. equi-trojani Asch. & Sint. numidica (de Lannoy ex Carrière) E. Murray,
ex Boiss., Fl. Orient. 5: 701. 1884; Abies alba Mill. Kalmia 12: 27. 1982. Type not designated. Fig. 13
subsp. equi-trojani (Asch. & Sint. ex Boiss.) Asch.
& Graebn., Syn. Mitteleurop. Fl. 1: 192. 1897; Abies
Etymology
nordmanniana (Steven) Spach var. equi-trojani
(Asch. & Sint. ex Boiss.) Guin. & Maire, Bull. Soc. The species epithet refers to Numidia, a Roman
Bot. France 55: 186. 1908; Abies equi-trojani (Asch. name for what is now NE Algeria.
& Sint. ex Boiss.) Mattf., Mitt. Deutsch. Dendrol.
Ges. 1925 (35): 29. 1925. Type: Turkey: [“M. Fola,
Vernacular names
in declivib. mont. Gargari”], P. E. E. Sintenis 523
(holotype not located, isotype K). Algerian fir; Sapin d’Algérie, Sapin des Babors (French)

Abies bornmuelleriana Mattf., Notizbl. Bot. Gart.

Description
Berlin-Dahlem 9: 239. 1925; Abies nordmanniana
(Steven) Spach subsp. bornmuelleriana (Mattf.) Trees to 20 m tall, d.b.h. to 0.5–0.8 m; trunk monopo-
Coode & Cullen, Notes Roy. Bot. Gard. Edinburgh dial, straight, columnar, terete, but on exposed sites
26: 167. 1965; Abies nordmanniana (Steven) Spach often twisted and forked; crown broad pyramidal,
var. bornmuelleriana (Mattf.) Silba, Phytologia 68: usually dense but irregular. Bark smooth and grey in
21. 1990. young trees, fissured and scaly, dark grey brown in
old trees. Branches of first order continuous to base
of trunk, spreading, the lower ones curved down;
Description
branches of second order dense, spreading horizon-
Vegetative buds (slightly) resinous; young foliage tally, ascending near the top of the tree. Branchlets
shoots (branchlets) glabrous. Cusps on bracts short, stout, firm, greyish brown or yellowish, or shining
without a distinct midrib. orange-brown, prominently ridged between the
leaves, glabrous; leaf scars circular. Vegetative buds
ovoid or broad conical, 5 × 4 mm, not resinous, or are Taxus baccata and Juniperus communis. At lower
only slightly resinous at base; bud scales ovate, ob- elevations broad-leaved trees, e.g. Acer obtusatum,
tuse or acute, appressed, brown, persisting several A. campestre, A. monspessulanum, Quercus spp. and
years. Leaves spirally arranged, radially spreading, Ilex aquifolium become more numerous; the only
especially on shoots in the periphery of the crown, broad-leaved species accompanying Abies and Ced-
those on lower shaded branches sometimes more or rus to the tree limit is Populus tremula. A common
less pectinate, on cone bearing shoots likewise radial, associated shrub is Genista numidica.
the upper leaves almost recurved, (1–)1.5–2(–2.5) cm
long, 2–3 mm wide, twisted or curved at base, linear
Conservation
or ligulate-linear, flattened, keeled by a prominent
midrib below, (dark) green, glaucous near the apex, Abies numidica has a very limited distribution in the 109
two whitish green bands below; apex obtuse or faint- Kabylie Range with an estimated area of occupancy
ly emarginate, rarely acutish on coning shoots. Sto- (AOO) below 20 km² in only two major populations.
mata especially near the apex or in a median groove The trees are legally protected against cutting but
above, in two bands divided by a midrib below. Pol- forest fires and other destructive events could well
len cones lateral, crowded, 1–2.5 cm long, yellowish. move the species into a higher category of threat.
Seed cones lateral, erect; with short, scaly peduncles; IUCN: VU (D2)
cylindric, with acutish, often papilliform apex, 12–18
cm long, 4–6 cm wide, light green tinged with purple
Uses
when immature, maturing to green with purple hue,
becoming light purplish brown when ripe; cone ra- Algerian fir is not exploited for timber. It is a suitable
chis persistent, narrowly conical, brown. Seed scales tree for cultivation, but it is sensitive to low tempera-
cuneate flabellate, length × width at mid-cone 2–3 × tures in winter and to air pollution in urban environ-
2.5–3.5 cm; surface smooth, slightly striated, puberu- ments. It is mostly cultivated in countries around the
lent on exposed parts; upper margin entire, repand; Mediterranean Sea, where it is sometimes planted in
base pedicellate. Bracts spathulate, with small cusps, hedges as it takes trimming well. Few cultivars are
0.8–1 cm long, included, rarely the cusps exserted known and the species is mostly grown from seed
in lowest part of cone. Seeds cuneate, 6–8 mm long, collected in situ; due to political circumstances in
light brown; seed wings cuneate-oblong, 12–15 mm Algeria it has been difficult to obtain seeds in recent
long, light brown, tinged with purple. decades. North of the Alps this species is still present
in botanic gardens and arboreta (e.g. Chèvre-loup
near Versailles, France), but it is becoming rare since
Distribution
dead trees are scarcely replaced with good stock
N Algeria (Kabylie Range, Mts. Babor and Tababor). from wild collected seed. Seeds used from planted
TDWG codes: 20 ALG trees are useless due to the ease with which this spe-
cies hybridizes with its European congeners.
Ecology
Abies numidica occurs in very isolated relict popula- Abies pindrow (Royle ex D. Don) Royle, Ill. Bot.
tions on north and east facing slopes of high moun- Himal. Mts. 1: t. 86. 1836.
tains in the Algerian Atlas. Its altitudinal range is
between 1800 m and 2000 m a.s.l. (Nitzelius, 1969),
Etymology
but Liu (1971) relates that occasional trees may
be found as low as 1220 m and as high as 2010 m. The species epithet takes up one of the vernacular
The soils are calcareous and rocky. The climate has names of this fir, as coined by Royle from its usage
warm, dry summers and cool, wet winters; the an- in the Simla Hills.
nual precipitation ranges between 1500 mm and
2000 mm, much of this falls as winter snow. The
Vernacular names
species may form pure stands, but more commonly
it is mixed with Cedrus atlantica, which is either co- Pindrow fir, West Himalayan fir; pindrau (Himach-
dominant or dominant. Other associated conifers al); badar (Kashmir); ragha (Kumaon)
 Description Ecology
Trees to 60 m tall, d.b.h. to 2.5–3 m; trunk mono­ Abies pindrow is a species of high mountains, occur-
po­dial, straight, columnar, terete; crown narrowly ring between 2000 m and 3300 m a.s.l. (occasion-
conical or pyramidal, often rather open and irregu- ally as high as 3700 m; Liu, 1971), on alpine lithosols.
lar. Bark of young trees smooth, light grey, turning The climate is cool, moist monsoon, with abundant
greyish brown, in old trees becoming longitudinally precipitation, but less than in the eastern Himalayas,
fissured, rough and scaly, dark grey brown. Branches much of it falling as snow. It occurs in pure stands
of first order relatively short, spreading horizontally; or in association with Picea smithiana, Pinus walli-
branches of second order similar. Branchlets stout, chiana, Tsuga dumosa and Cedrus deodara; at lower
110 firm, light greyish pink, or light brown, soon be- elevations broad-leaved trees, e.g. Quercus seme-
coming grey, faintly ridged and grooved, glabrous; carpifolia, Q. dilatata, Juglans regia, Aesculus indica,
leaf scars circular. Vegetative buds ovoid globose, Acer spp., Prunus spp., and Ulmus spp. become more
6–8 mm long, often very resinous, but most resin important, replacing the conifers below 1600 m.
has disappeared after one growing season; bud
scales broad triangular, with laciniate or erose mar-
Uses
gins, purplish brown or dull brown, persisting sev-
eral years. Leaves spirally arranged, pectinate in two Pindrow fir is an important timber tree in the
opposite level rows or standing out more radially, Himalayas, where its timber is used in construc-
the uppermost leaves shorter and directed obliquely tion (house building), in particular for interior work
forward, on coning shoots assurgent, (2.5–)3–6(–7) such as floor boards, ceilings, and stairs. In some
cm long, 1.3–2 mm wide, twisted at base, narrowly parts shingles are used for roofing. Another applica-
linear, flattened, grooved above, lustrous (dark) tion of its wood is for fruit cases and tea boxes. This
green above, two greenish white bands below; apex species remains uncommon in cultivation in Europe
sharply bifid or emarginate, or obtuse (acute on and is regularly misidentified, with trees named
coning shoots. Stomata in two narrow bands, sepa- A. pindrow var. intermedia turning out to belong to
rated by a midrib below, none or a few near apex A. spectabilis (Rushforth, 1987). It requires a mildly
above. Pollen cones lateral, crowded on the under- cool and wet climate, such as prevails in the western
side of shoots, 1–1.5 cm long, yellowish brown. Seed parts of the British Isles.
cones lateral, erect, short pedunculate, cylindrical or
ovoid-­cylindrical, with obtuse apex, 10–14 cm long, 2 varieties are recognized:
5–6(–7) cm wide, violet blue when immature, matur-
ing to purplish brown and becoming dark reddish Abies pindrow (Royle ex D. Don) Royle var.
brown when ripe; cone rachis persistent, narrowly pindrow. Pinus pindrow Royle ex D. Don, London
conical, dark brown. Seed scales flabellate, length × Edinburgh Philos. Mag. & J. Sci. 8: 255. 1836; Abies
width at mid-cone 2.5–3.5 × 3–4 cm; surface smooth, webbiana (Wall. ex D. Don) Lindl. var. pindrow
slightly striated, puberulent on exposed parts; upper (Royle ex D. Don) Brandis, Forest Fl. NW India:
margin entire, slightly incurved; base pedicellate. 528. 1874. Type: Illustration in Royle, Ill. Bot.
Bracts obovate or semi-orbicular, with small cusps, Himal. Mts. 1: t. 86. May 1836 (lectotype, vide
1 cm long, entirely included. Seeds cuneate, angu- D. Don in Lambert, 1837).
lar, 10–12 mm long, shining dark brown or blackish;
seed wings cuneate, 15–20 mm long, brown. Description
Leaves 3–6(–7) cm long, on vegetative shoots pec-
Distribution tinately arranged; apex sharply bifid or emarginate;
stomata only in two bands on the underside.
Himalaya: from Afghanistan east to Nepal, Kara­
koram Range.
TDWG codes: 34 AFG 40 NEP PAK WHM-HP WHM- Distribution
JK WHM-UT
Himalaya: east to Nepal, Karakoram Range.
TDWG codes: 34 AFG 40 NEP PAK WHM-HP WHM-
JK WHM-UT
 nar and terete in sheltered trees, but often twisted
Conservation
and forked; crown in young trees narrowly conical,
IUCN: LC old trees irregular, open or dense. Bark of young
trees smooth, dark grey, in old trees rough and scaly.
Abies pindrow (Royle ex D. Don) Royle var. Branches of first order long, curved downward, the
brevifolia Dallim. & A. B. Jacks., Handb. Conif.: ones near the top ascending; branches of second
126. 1923. Type not designated. order dense, spreading horizontally and ascending.
Branchlets stout, very firm, reddish brown or green-
Abies gamblei Hickel, Bull. Soc. Dendrol. France 70: ish brown, turning grey, faintly ridged between the
38. 1929; Abies pindrow (Royle ex D. Don) Royle sub- leaves, glabrous; leaf scars circular or angular, large,
sp. gamblei (Hickel) Rushforth, Int. Dendrol. Yearb. purplish grey. Vegetative buds ovoid-globose, 5–6 × 111
1998: 63. 1999. 4–4.5 mm, not resinous to very resinous; bud scales
triangular, keeled and free at the apices, light reddish
brown or purplish brown. Leaves spirally arranged,
Description
spreading radially and perpendicularly from shoots,
Leaves 2.5–3.8 cm long, more or less radially ar- or more or less pectinate, the upper leaves often
ranged (not pectinate); apex obtuse on vegetative recurved, in shaded shoots the lower leaves some-
shoots; sometimes also a few stomata on the upper- what pectinate, 0.6–2 cm long, 2–3 mm wide, not or
side. only slightly twisted at base, linear-­ligulate, carinate
or slightly flattened, rigid, grey-green or glaucous
green; apex obtuse, acute or acuminate. Stomata
Distribution
above in several rows, below in two bands separated
India: Garhwal and Kashmir Himalayas (Chamba by a midrib and bordered by broad margins. Pol-
District). len cones lateral, crowded, 0.5–0.7 cm long, yellow-
TDWG codes: 40 WHM-HP WHM-JK ish with red or violet microsporophylls. Seed cones
lateral, erect, short pedunculate, cylindrical, with
obtuse, often papilliform apex, 9–14(–18) cm long,
Conservation
3–4(–5) cm wide, greenish purple when immature,
IUCN: LC becoming dark or light brown when ripe; cone ra-
chis persistent, narrowly conical, purplish brown.
Seed scales cyathiform or cuneate-flabellate, length
Abies pinsapo Boiss., Not. Abies Pinsapo: 8. 1838 × width at mid-cone 2.5–2.8 × 2.2–2.5 cm; surface
[& Biblioth. Universelle Genève 13: 406. 1838]. Fig. smooth, slightly striated, yellowish pubescent on ex-
14, 15 posed parts; upper margin entire, somewhat undu-
late, incurved; base long pedicellate. Bracts oblong;
apex obcordate, with a tiny cusp, 1–1.3 cm long,
Etymology
entirely included. Seeds cuneate-oblong, 8–10 mm
The species epithet ‘pinsapo’ is the Andalucian name long, light brown; seed wings cuneate-oblong, 13–20
for this fir. mm long, light brown.

Vernacular names Distribution

Spanish fir (including Moroccan fir); pinsapo (Span- N Morocco, S Spain.
ish) TDWG codes: 12 SPA-SP 20 MOR-MO

Description Ecology
Trees to 30 m tall, but most trees in nature smaller, Abies pinsapo is a species of the north slopes of high
d.b.h. to 1–1.5 m; trunk monopodial, straight, colum- mountains, where it occurs between 1100 m and
 2000 m a.s.l. (var. pinsapo in S Spain) and 1400 m Abies pinsapo Boiss. var. pinsapo. Type: Spain:
and 2100 m a.s.l. (the Moroccan variety). It grows on Malaga, Sierra Bermeja, N of Estepona [“Sierra de
rocky soils derived from dolomitic limestone or ser- la Nieve”], E. Boissier s.n. (holotype not located,
pentine, with deep drainage. The climate is montane, isotype K).
with a mediterranean influence: dry, warm summers
alternate with cool, moist winters, with annual pre-
Description
cipitation around 1000 mm. Both varieties occur in
pure, scattered stands (very rare in Spain) or mixed Leaves spreading radially and perpendicularly from
with Cedrus atlantica (Morocco) or Pinus pinaster shoots; apex obtuse or acute; resin ducts medial.
(Spain). Commonly, broad-leaved trees, e.g, Quer- Seed cones 9–14 cm long, 3–4 cm wide.
112 cus ilex, Q. lusitanica, and Q. canariensis (Morocco),
and (often sclerophyllous) shrubs, e.g. Ulex balticus,
Distribution
Cistus spp., Pistacia lentiscus, Daphne laureola and
Berberis hispanica are mixed with scattered A. pin- S Spain: Prov. Granada, Málaga.
sapo at lower elevations. TDWG codes: 12 SPA-SP

Uses Conservation
Spanish fir (including its Moroccan variety) is no Abies pinsapo var. pinsapo is restricted to a small
longer exploited for its timber. This species is one of number of disjunct populations, the largest one situ-
the most attractive firs in horticulture with its rigid ated within the Sierra de las Nieves National Park.
needles spreading all around the shoots and espe- Fires, some started by tourists, are a hazard threat-
cially some more glaucous-leaved trees have been ening especially some of the smaller, isolated stands
selected as cultivars and are widely planted. While with destruction. Within and without the National
most remaining trees at high altitude in southern Park grazing by goats, a pan-Mediterranean men-
Spain are short and slow growing, Spanish fir can ace, hampers natural regeneration. Erosion of the
grow tall in less exposed sites with ample moisture. already thin and depleted soil adds to the low ability
For garden purposes slower growing ‘dwarf ’ forms to establish new trees naturally. A planting scheme
are more popular and a few cultivars, some derived to restore depleted populations is being implement-
from witches brooms found in natural populations, ed.
meet these requirements. Apparently this species IUCN: VU (D2)
can be easily hybridized with a number of other
species, e.g. A. alba, A. cephalonica, A. numidica (its Abies pinsapo Boiss. var. marocana (Trab.) Cebal-
closest relative), A. nordmanniana and A. pindrow; los & Bolaño, Bol. Inst. Nac. Invest. Agron. 1 (2): 18.
some of these crosses have been the parental stock 1928. Abies marocana Trab., Bull. Soc. Bot. France
of further cultivars. Many hybrids resulting from 53: 154. 1906. Type not designated.
(controlled or spontaneous) crossings in cultiva-
tion have been given nothospecific names e.g. Abies Abies tazaotana S. Côzar ex Villar, Types Sols Afri-
×vilmorinii Matf. for the cross between A. pinsapo que N. 1: 80. 1947; Abies pinsapo Boiss. var. tazaotana
and A. cephalonica. Since seed produced from these (S. Côzar ex Villar) Pourtet, Ann. Ecole Natl. Eaux 9
F1 crossings in cultivation is likely to have been fer- (1): 100. 1954; Abies pinsapo Boiss. subsp. tazaotana
tilized by unknown fir pollen, perpetuation of the (S. Côzar ex Villar) R. Govaerts, World Checklist
hybrid is only guaranteed by vegetative propagation Seed Pl. 1 (1): 6. 1995.
or by renewed controlled crossing of the parent spe-
cies. Such plants should receive cultivar names, not
Description
taxon names.
Leaves more ore less pectinately arranged, especially
2 varieties are recognized: on lower branches, acute to acuminate; resin canals
marginal. Seed cones 10–18 cm long, 3.5–5 cm wide.
 in two narrow bands separated by a midrib below.
Distribution
Pollen cones lateral, pedunculate, solitary or a few
N Morocco (Rif Mts.). together in leaf axils, 1.5–2.5 cm long, yellow with
TDWG codes: 20 MOR-MO red microsporophylls. Seed cones lateral, erect,
short pedunculate, cylindric-conical, with obtuse
or truncate apex, 15–20 cm long (–30 cm on culti-
Conservation
vated trees), 5–8 cm wide, green and often tinged
IUCN: NT with red when immature, maturing to greenish grey,
with yellowish bracts, ripening to light grey brown
with light, orange brown bracts; cone rachis persist-
Abies procera Rehd., Rhodora 42: 522. 1940. Type: ent, narrowly conical, purplish brown. Seed scales 113
USA: Washington, Columbia River, D. Douglas s.n. cuneate-flabellate or cyathiform, length × width at
(holotype not located, isotype K). Fig. 16 mid-cone 2.5–3 × 2.5–3.5 cm; surface smooth, pu-
bescent to hirsute, with yellowish grey hairs; upper
margin entire, strongly incurved; base long pedicel-
Etymology
late. Bracts spathulate-obcordate, with laciniate up-
The species epithet means ‘slender’ or ‘tall’ and re- per margins and a long, curved cusp, 3–4 cm long,
fers to the shape of the trees in dense forest stands. exserted, reflexed, covering much of the exposed
parts of the seed scales. Seeds cuneate-oblong, 12–15
mm long, pale reddish brown; seed wings obovate
Vernacular names
or cuneate-oblong, ca. 20 × 15 mm, straw coloured.
Noble fir
Distribution
Description
USA: NW California, Oregon, Washington (Cas-
Trees to 70–80 m tall, d.b.h. to 2.5–3 m; trunk mo- cade Range, parts of Coast Range).
nopodial, straight, columnar, terete, often bare of TDWG codes: 73 ORE WAS 76 CAL
branches to a considerable height and very regular;
crown narrowly pyramidal or more conical, in forest
Ecology
stands often only ⅓ or less of total tree height. Bark
of young trees smooth, thin, with resin blisters, light A magnificent tree, occurring from the foothills of
grey with a purplish tinge, in old trees rather smooth, mountains in W Washington to high mountain sides
except on lower half of trunk where it breaks into ir- in Oregon, between 60 m and 2700 m a.s.l. It is most
regular plates, greyish brown. Branches of first order abundant in the mountains of the Cascade Range,
spreading horizontally, the lower pendulous, but of- on a variety of mountain soils with ample moisture
ten contorted in old trees; branches of second order available to the vegetation. The climate is cool tem-
spreading horizontally and ascending near the ends perate, with short summers and snowy winters, the
of main branches. Branchlets slender, firm, reddish annual precipitation ranging from 1750 mm to 2600
brown in the first year, soon purplish brown, faintly mm, much of it as snow. It mainly grows in the Ca-
ridged and grooved, pubescent, but almost hidden nadian Life Zone, but also in the lower Transition
by leaves; leaf scars oval or obovate. Vegetative buds Zone, where it can be associated with several other
ovoid-globose, hidden by leaves, 3 × 2 mm, slightly conifers, e.g. Tsuga heterophylla, Picea sitchensis and
resinous; bud scales ovate, dark purplish red, persist- Thuja plicata near the coast, Pseudotsuga menziesii,
ing several years. Leaves spirally arranged, strongly Abies grandis, Pinus spp. in much of its range, and
assurgent on all shoots, the lower leaves spreading Abies lasiocarpa, A. amabilis, Tsuga mertensiana,
and S-curved, 1–2.5(–3.5) cm long, those midway Picea engelmannii, Larix occidentalis at higher el-
of a year’s shoot longest, 1.5–2 mm wide, curved at evations. Common shrubs are Rhododendron spp.,
base, basi-falcate or falcate-linear, flattened or cari- Vaccinium spp. and Ribes spp. Abies procera can be
nate, glaucous green above, glaucous white bands dominant, but occurs rarely in pure stands (Fowells,
below; apex obtuse (rarely notched), acute on con- 1965).
ing shoots. Stomata in several to many rows above,
 lowish brown, light brown or pink-brown, later grey,
Conservation
smooth or slightly pubescent, shining and glabrous,
IUCN: LC ridged between the leaves; leaf scars circular, light
grey. Vegetative buds ovoid or broad conical, 5–8 ×
4–6 mm, resinous, but resin soon disappearing; bud
Uses
scales triangular, keeled, light brown, persistent for
Noble fir attains large dimensions and grows an several years. Leaves spirally arranged, spreading ra-
extremely straight bole under favourable site con- dially, more or less pectinate below, strongly assur-
ditions. Almost pure natural stands can yield large gent to recurved above, especially on cone bearing
volumes of timber per ha. Its wood is of higher qual- shoots and on leading shoots higher in the crown,
114 ity than that of other firs in North America due to (1.2–)1.5–3.5 cm long, 1.9–2.5 mm wide, twisted
greater strength and indeed its size. Besides general or recurved at base, linear, but the shortest leaves
construction and carpentry applications, special oblanceolate, flattened, longitudinally grooved
uses have been propellors of airplanes and ladders, above, glossy green above, 2 greenish white bands
now mostly replaced by various metals. The odour- below; apex emarginate, obtuse or acute. Stomata in
less, white wood is excellent for making boxes. two bands separated by a midrib below, occasionally
Young trees make attractive Christmas trees with a few near apex above. Pollen cones lateral, pendant,
their dense, upturned glaucous leaves. In amenity 1–1.5 cm long, yellow, with reddish microsporo-
planting and horticulture this fir is one of the more phylls. Seed cones lateral, erect, often clustered, short
popular and commonly used species and several pedunculate, ovoid or ovoid-oblong, with obtuse or
cultivars are known. It was introduced to England papilliform, sometimes umbilicate apex, 5–8(–9) cm
by David Douglas in 1830. It has the largest seed long, 2.5–3.5 cm wide, bluish purple when imma-
cones of all species, with attractive, exserted yellow- ture, maturing to blue-grey, ripening to grey-brown;
ish bracts. It is unsuitable in climates with summer cone rachis persistent, thick, cylindro-conical to
droughts or less than a good supply of rain spread fusiform, purplish brown or light brown. Seed scales
evenly in the year as its bark tends to split open dur- broadly flabellate to cyathiform, length × width at
ing dry spells. mid-cone 1.5–2 × 2–2.5 cm; surface smooth, puberu-
lous; upper margin thin, entire, erose or irregularly
fissured; base pedicellate. Bracts spathulate, with a
Abies recurvata Mast., J. Linn. Soc., Bot. 37: 423. short cusp, 1–1.5 cm long, entirely included, or occa-
1906. sionally the cusps exserted. Seeds cuneate, 5–8 × 3–4
mm, light brown; seed wings broadly cuneate, 6–10
× 5–6 mm, violet-blue, turning brown.
Etymology
The species epithet refers to the often recurved or
Distribution
reflexed leaves (needles) on the shoots.
China: SW Gansu, Sichuan, NW Yunnan (?), SE Xi-
zang [Tibet].
Vernacular names
TDWG codes: 36 CHC-SC CHN-GS CHT
Min fir; min kiang lien sha (Chinese)
Ecology
Description
Min fir (both varieties) is a high mountain species of
Trees to 40–60 m tall, d.b.h. to 1.5–2.5 m; trunk mo- SW China, occurring between 2300 m and 3600 m
nopodial, straight, columnar, terete; crown conical a.s.l. or even higher. It grows usually on grey-brown
or pyramidal, flat topped in old trees. Bark of young mountain podzols. The climate is cold, moist, with
trees smooth, soon with papery flakes, grey or pink annual precipitation between 700 mm and 1000
brown, in old trees rough, flaky, breaking into small mm. Both varieties are usually constituents of a
scales, dark grey brown. Branches of first order rela- mixed coniferous forest type, with among other spe-
tively short, horizontally spreading; branches of cies A. squamata, Picea likiangensis var. rubescens,
second order similar. Branchlets slender, firm, yel- P. asperata, and Larix potaninii; Picea purpurea and
Abies fargesii var. faxoniana are mainly found with often recurved at base; apex obtuse or acute; resin
the ‘typical’ variety, and A. fabri with var. ernestii. ducts medial. Seed cones with obtuse or papilliform
Betula albosinensis is the only common broad-leaved apex; rachis usually cylindro-conical.
tree at higher elevations, but lower down the slopes
other genera, e.g. Acer, Populus, but also different
Distribution
conifer species, e.g. Tsuga chinensis, Picea brachytyla
var. complanata and Pinus armandii become more China: SW Gansu, Sichuan.
abundant. TDWG codes: 36 CHC-SC CHN-GS

Uses Conservation
115
A timber tree in western China, heavily exploited This variety is limited to an area along the Min Riv-
until recently when the Chinese government finally er, south of Songpan in N Sichuan, where it may not
decided to preserve its remaining old growth forests occupy more than 20 km².
in the western provinces. Its timber was used mainly IUCN: VU (D2)
for construction and carpentry work. The type col-
lection (of var. recurvata) was collected by Ernest Abies recurvata Mast. var. ernestii (Rehd.)
H. Wilson on his first expedition to western China C. T. Kuan, Notes Roy. Bot. Gard. Edinburgh 41:
in 1903; the species was introduced to horticulture 536. 1984. Abies ernestii Rehd., J. Arnold Arbor.
in the USA and UK from seed collected by him on 20: 85. 1939; Abies chensiensis Tiegh. var. ernestii
subsequent journeys to the Min River drainage. As (Rehd.) T. S. Liu, Monogr. Gen. Abies: 135. 1971.
with most Chinese species in Abies, it remains a Type: China: Sichuan, Daxue Shan, Kangding, [“NE
dendrological collector’s item and has not entered of Tachien-lu, Tapao-shan”], E. H. Wilson 2090
the common gardening trade. A main reason for (holotype A). Fig. 17
this is undoubtedly the unavailability of seed from
its country of origin for a long period after the ef-
Description
forts of the early 20th century’s plant collectors came
to an end. Renewed collecting, made possible in the Buds ca. 8 × 6 mm; shoots yellowish brown, faint-
last few decades in partnership with Chinese bota- ly pubescent. Leaves 2–3.5 cm long, more or less
nists, has been undertaken under more restricting straight; apex emarginate on vegetative shoots; resin
conditions and the results have largely remained ducts marginal. Seed cones often with an umbilicate
within the confines of major botanic gardens. Even apex and fusiform rachis.
if trees in cultivation produce viable seed, unless
they are grown in complete isolation from other
Taxonomic notes
species of Abies, that seed is likely to produce plants
with a mixture of genes from almost any of those In Flora of China 4: 51 (1999) this variety is treated
other species. as a distinct species, while within A. recurvata no
infraspecific taxa are recognized.
2 varieties are recognized:
Distribution
Abies recurvata Mast. var. recurvata. Type: China:
Sichuan, Min River, Songpan, [“south of Sung Pan, China: SW Gansu, W Sichuan, NW Yunnan (?), SE
Min Valley”], E. H. Wilson 3021 (holotype K). Xizang [Tibet].
TDWG codes: 36 CHC-SC CHN-GS CHT
Description
Conservation
Buds ca. 5 × 4 mm; shoots usually light brown or
pink-brown, glabrous. Leaves (1.2–)1,5–3 cm long, IUCN: LC
 Abies religiosa (Kunth) Schltdl. & Cham., Linnaea ture, becoming dark (purplish) brown with brown
5: 77. 1830. Pinus religiosa Kunth, in Humboldt bracts when ripe; cone rachis persistent, narrowly
et al., Nov. Gen. Sp. Pl. 2 (5): 5. 1817. Type: conical, dark brown. Seed scales cuneate-flabellate,
Mexico: Guerrero, F. W. H. A. von Humboldt & length × width at mid-cone 2–3 × 3–3.5 cm; surface
A. J. A. Bonpland s.n. (holotype P). smooth, puberulent; upper margin rounded, entire;
base pedicellate. Bracts linear-spathulate, with a
Abies colimensis Rushforth & Narave, Notes Roy. ­tapering cusp, 3–3.5 cm long, exserted, reflexed, cov-
Bot. Gard. Edinburgh 46: 105. 1989. ering part of the seed scales. Seeds cuneate-oblong,
10 × 5 mm, shining brown; seed wings cuneate-­
dolabriform, 10–15 mm long, brown.
Etymology
116
The species epithet refers to the traditional religious
Distribution
significance of this tree to the peoples of Guatemala
and Mexico. W Guatemala; Central and S Mexico (highlands).
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
PU MXC-TL MXE-GU MXE-HI MXE-NL MXE-QU
Vernacular names
MXE-SL MXE-TA MXG-VC MXN-SI MXS-GR MXS-MI
Sacred fir MXS-JA MXS-OA MXT-CI 80 GUA

Description Ecology
Trees to 50–60 m tall, but usually not exceeding 40 Abies religiosa is a high mountain species, occurring
m, d.b.h. to 1.5–2 m; trunk monopodial, straight, between 1200 m and 4100 m a.s.l., but more com-
columnar, terete; crown pyramidal or conical. Bark monly between 2100 m and 3100 m, usually on well
of young trees smooth, greyish white, of old trees drained mountain soils of volcanic origin. The cli-
rough, deeply fissured, breaking into small plates, mate is cool, moist oceanic on ranges near the coast,
grey brown. Branches of first order long, slender, colder with more snow in the interior, with abun-
ascending, later becoming horizontal, the lowest dant precipitation. There are pure stands of this fir
pendant; branches of second order spreading hori- at the higher elevations, but it is often mixed with
zontally or slightly ascending. Branchlets slender, Pinus montezumae, P. hartwegii, in the north of its
especially the lower vegetative shoots, firm, reddish range also with Pseudotsuga menziesii var. glauca;
brown to purple red, ridged and grooved between at lower elevations Quercus spp., Alnus acuminata,
the leaves, slightly pubescent in the grooves or gla- Prunus serotina, and Arbutus spp. become more
brous; leaf scars circular, light grey. Vegetative buds abundant. Shrubs are e.g. Vaccinium spp., Andro­
ovoid globular, small, covered with resin; bud scales meda spp., Ribes spp., and Fuchsia spp.
broadly ovate, keeled, reddish brown, persisting
several years. Leaves spirally arranged, pectinate on
Conservation
vegetative shoots, directed slightly forward, on con-
ing shoots more or less radial and slightly assurgent, IUCN: LC
(1–)1.5–3(–3.5) cm long, 1.2–1.6 mm wide, twisted
at base, narrowly linear, grooved above, flattened,
Uses
shining dark green, or occasionally more glaucous,
with two whitish bands below; apex acute or obtuse- In Guatemala and Mexico the timber of this species
truncate. Stomata absent or only a few above, in two is used for light indoor construction and general
narrow bands separated by a midrib below. Pollen carpentry. Wholesale logging is unlikely to occur
cones lateral, more or less pendulous, 1–1.5 cm long, because this tree has traditional religious signifi-
yellow, with red microsporophylls. Seed cones later- cance to Native Americans. With the conversion to
al, sometimes subterminal, usually erect, with short, (Roman Catholic) Christianity and hispanisation of
often curved peduncles, ovoid oblong, cylindrical or the populations of this part of Latin America these
curved, with obtuse apex, (8–)10–16 cm long, 4–6 cm traditions were incorporated into the new modes of
wide, violet blue with yellowish bracts when imma- worship and at times of religious festivals churches
are being decorated with the foliage of this fir. This below. Pollen cones lateral, densely clustered on
species also has a very special ecological significance the underside of shoots, 1 cm long, yellowish, with
since it serves as the hibernation tree (in a very lim- red microsporophylls. Seed cones lateral, erect, of-
ited part of its range in south-central Mexico) for ten crowded, almost sessile, ellipsoid-cylindrical,
many millions of Monarch butterflies (Danaus plex- with obtuse or acutish apex, 5–8 cm long, 2–3 cm
ippus) yearly making one of the most spectacular wide, light or dark purple with greenish or red-
displays in all of nature. No one, despite decades of dish bracts when immature, becoming dark brown
research into the phenomenon, yet knows why these with brown bracts when ripe; cone rachis persist-
butterflies choose those trees in that particular lo- ent, narrowly conical, blackish purple. Seed scales
cality to congregate in such numbers, arriving from reniform, length × width at mid-cone 1 × 1.6 cm;
as far away as E Canada. surface smooth, pubescent on the exposed part; up- 117
per margin entire, incurved; base pedicellate. Bracts
obcordate-quadrangular, 1–1.3 cm long, included or
Abies sachalinensis (F. Schmidt) Mast., Gard. exserted, recurved, the upper bracts often included
Chron., ser. 2, 12: 588. 1879. in ripe cones. Seeds cuneate, 6 mm long, lustrous
brown with black spots; seed wings broad cuneate,
5 × 5 mm, purplish black.
Etymology
The species epithet refers to the island of Sakhalin
Distribution
from where its was first described.
Japan: Hokkaido; Russian Far East: southern Kuril
Islands, Sakhalin.
Vernacular names
TDWG codes: 31 KAM KUR SAK 38 JAP-HK
Sakhalin fir; akatodo, todo-matsu (Japanese); Pikhta
sakhalinskaya (Russian)
Ecology
Sakhalin fir and its varieties occur from near sea
Description
level on the coast to an elevation of 1650 m a.s.l. in
Trees to 25–30 m tall, d.b.h. to 0.8–1 m; trunk mo- the mountains. The soils are well drained but moist
nopodial, straight, columnar, terete; crown broad throughout the year, due to abundant precipitation
pyramidal, often flat topped in old trees; dead in a cool to cold, maritime climate. In the north of
branches remain on the bole. Bark of young trees its range the species is more common at elevations
smooth, with resin blisters, grey with a brown tinge, between 800 m and 1100 m, where it is mixed with
of old trees in lower part of the bole breaking into Picea jezoensis, P. glehnii, Larix gmelinii var. japonica
irregular plates, grey brown. Branches of first or- or Pinus pumila at the highest limit of trees. At lower
der long, slender, mostly ascending, the lower ones elevations pure stands occur, below 800 m broad-
spreading horizontally; branches of second order leaved-trees, e.g. Betula ermanii, Acer spp., Quercus
similar. Branchlets slender, firm, red-brown or mongolica var. grossesserata, Castanea crenata, Kalo-
brown, sometimes grey-brown, with faint ridges panax septemlobus, and Magnolia hypoleuca become
and shallow grooves, minutely pubescent in grooves more abundant.
with red-brown hairs; leaf scars circular, small. Veg-
etative buds ovoid, small, very resinous; bud scales
Uses
reddish brown, persisting several years. Leaves spi-
rally arranged, spreading more or less radially, the This species is mainly logged for the manufacture
upper leaves directed forward, covering shoot, on of wood pulp used in the paper industry; its tim-
coning shoots slightly assurgent, 1.2–3.5 cm long, ber is of low quality for construction and carpentry.
1–1.2 mm wide, (strongly) twisted or curved at base, As an amenity tree it is little used outside the cool
linear, grooved above, flattened, glossy dark green to cold maritime climate of northern Japan and the
above, two greenish white bands below; apex emar- Russian Far East. It is in cultivation in botanic gar-
ginate or obtuse. Stomata none or a few near apex dens and arboreta in Russia, northern Europe and
above, in two narrow bands separated by a midrib New England, USA, but rarely survives to maturity
 in countries with mild winters, where it will not go
Taxonomic notes
into prolonged winter dormancy and is susceptible
to spring frosts. Morphologically, this taxon is most similar to Abies
sachalinensis and on these grounds alone does not
4 varieties are recognized, one tentatively: seem to merit recognition as a distinct species. Its
status as a species is accepted by some Russian bota-
Abies sachalinensis (F. Schmidt) Mast. var. sacha- nists but disputed by others; to solve this problem
linensis. Abies veitchii Lindl. var. sachalinensis analysis of its DNA in comparison with several re-
F. Schmidt, Mém. Acad. Imp. Sci. Saint-Péters- lated eastern Asian firs is in preparation (Farjon &
bourg, sér. 7, 12 (2): 175. 1868; Abies nephrolepis Xiang unpublished data, Aug 2009).
118 (Trautv. ex Maxim.) Maxim. subsp. sachalinensis
(F. Schmidt) V. N. Voroshilov, Bjull. Moskovsk.
Distribution
Obsc. Isp. Prir., Otd. Biol. 96 (1): 133. 1991. Type:
Russia: Russian Far East, Sakhalin, 1862, F. Schmidt Russian Far East: Kamchatka (Kronotzky Bay).
s.n. (holotype not located, isotype K). TDWG codes: 31 KAM

Abies sachalinensis (F. Schmidt) Mast. var. corticosa

Ecology
Tatew., Trans. Sapporo Nat. Hist. Soc. 45: 70. 1935;
Abies sachalinensis (F. Schmidt) Mast. f. corticosa The occurrence of a small, isolated ‘grove’ of ca.
(Tatew.) Hayashi, Taxon. Phytogr. Japon. Conif.: 26. 30,000(?) trees occupying an area of no more than
1960. 20 ha in coastal lowland in the SE part of the pe-
ninsula of Kamchatka is something of an enigma.
There is some expression of doubt to its indigenous
Description
status in the Russian literature but there seems to be
Bark grey or brownish grey. Leaves with medial res- no thorough scientific research published under-
in ducts. Seed cones dark purple when immature; pinning its status either as an introduction (by na-
bracts exserted, reddish turning brown. tive people?) or as a ‘Tertiary relict’ as some others
believe. A scientific assessment addressing its indi-
genity is urgently needed, because its locality seems
Distribution
totally at odds with what is known of the biogeo­
Japan: Hokkaido; Russian Far East: Kuril Is., Sakha- graphy of its nearest relatives (Sect. Balsamea) in the
lin. genus Abies.
TDWG codes: 31 KUR SAK 38 JAP-HK
Conservation
Conservation
IUCN: VU (D2)
IUCN: LC
Abies sachalinensis (F. Schmidt) Mast. var. mayri-
Abies sachalinensis (F. Schmidt) Mast. var. gracilis ana Miyabe & Kudô, Trans. Sapporo Nat. Hist.
(Kom.) Farjon, Pinaceae (Regnum Veg. 121): 83. Soc. 7: 131. 1919. Abies mayriana (Miyabe & Kudô)
1990. Abies gracilis Kom., Trudy Imp. S.‑Peter- Miyabe & Kudô, Icon. Ess. Forest Trees Hokkaido 1:
burgsk. Bot. Sada 20: 203. 1901; Abies sibirica Ledeb. 9, t. 3, 4. 1920. Type not designated.
var. gracilis (Kom.) Patschke, Bot. Jahrb. Syst. 48:
684. 1913. Type not designated.
Description
Bark smooth, whitish grey. Leaves with medial resin
Description
ducts. Seed cones light purplish, with greenish, ex-
Bark grey. Leaves with marginal resin ducts. Seed serted bracts turning light brown at maturity.
cones purple when immature; bracts exserted, turn-
ing brown at maturity.
 pendant; branches of second order dense, spread-
Distribution
ing out laterally or assurgent. Branchlets slender,
Japan: Hokkaido; Russian Far East: Sakhalin. firm, pale yellowish grey to fawn brown or yellowish
TDWG codes: 31 SAK 38 JAP-HK brown, soon grey, smooth or only faintly ridged or
more prominently ridged and grooved, with short
pubescence, soon glabrous; leaf scars small, circu-
Conservation
lar. Vegetative buds globose, 2–3 mm long, more or
IUCN: LC less resinous; bud scales reddish or yellowish brown,
persisting several years. Leaves spirally arranged,
Abies sachalinensis (F. Schmidt) Mast. var. nemo- the upper leaves covering shoot or assurgent, the
rensis Mayr, Monogr. Abiet. Japan. Reich.: 42, t. 3, f. lower leaves pectinate in two lateral planes, on con- 119
6. 1890. Abies nemorensis (Mayr) Miyabe & Kudô, ing shoots all leaves assurgent, (1–)1.5–3 cm long,
in Miyabe & Miyake, Fl. Saghalin: 745. 1915. Type 1.3–1.6 mm wide, twisted or curved at base, linear,
not designated. flattened, light green, with 2 greyish green bands be-
low; apex emarginate, obtuse, or acute with a callous
tip especially on coning shoots. Stomata occasion-
Description
ally a few near apex above, in two bands separated
Bark grey. Leaves with medial resin ducts. Bracts by a midrib below. Pollen cones lateral, crowded on
of seed cones included or only the small cusps ex- the underside of shoots, 1.5 cm long, yellow, with red
serted. microsporophylls. Seed cones lateral, erect, nearly
sessile, ovoid-cylindrical, with obtuse apex, 5–7.5 cm
long, 2.5–3.5 cm wide, violet blue when immature,
Distribution
maturing to bluish brown, or yellowish brown, be-
Japan: Hokkaido; Russian Far East: Sakhalin. coming (light) cinnamon or greenish brown when
TDWG codes: 31 SAK 38 JAP-HK ripe; cone rachis persistent, narrowly conical, dark
brownish purple. Seed scales broad flabellate, up-
per scales more cuneate, length × width at mid-cone
Conservation
1.5–1.7 × 2–2.2 cm; surface smooth, puberulent; up-
IUCN: LC per margin entire or serrulate-erose; base pedicel-
late. Bracts short, rounded, with a tiny cusp, 0.8 cm
long, entirely included. Seeds cuneate-oblong, 5–6
Abies sibirica Ledeb., Fl. Altaica 4: 202. 1833. mm long, brown; seed wings cuneate-oblong, 10–12
mm long, light brown.
Etymology
Distribution
The species epithet refers to its occurrence in Sibe-
ria. Across N Russia and Siberia, from Archangel’sk
eastward to the Amur River, southward to the
mountains along the Sino-Russian border and the
Vernacular names
Tien Shan Range.
Siberian fir; Pikhta sibirskaya (Russian); Xinjiang TDWG codes: 14 RUE RUN 30 ALT BRY CTA IRK
lengshan (Chinese) KRA TVA WSB YAK 31 AMU KHA 32 KAZ KGZ 36 CHX

Description Ecology
Trees to 35–40 m tall, d.b.h. to 1 m; trunk mono­podial, The ‘typical’ subspecies of Siberian fir is widespread
straight, columnar, terete; crown narrowly pyrami- across the Siberian taiga, where it occurs from near
dal or conical. Bark of young trees smooth, with sea level on the northern plains to 2000 m a.s.l. in
numerous resin blisters, grey or grey brown, finally the mountains. It remains well south of the arctic
breaking into plates in old trees. Branches of first or- tree limit in Siberia, in fact it is more common in
der short, slender, spreading horizontally, the lower W Siberia and the Altai Mountains, which have a
 less severe climate. The soils are usually of alluvial Abies sibirica Ledeb. subsp. semenovii (B. Fedtsch.)
origin, podzolic, and in the mountains also calcare- Farjon, Pinaceae (Regnum Veg. 121): 81. 1990. Abies
ous, well drained and free of permafrost. The climate semenovii B. Fedtsch., Bot. Centralbl. 73 (7): 210.
is cold continental, but not extreme in most parts 1898; Abies sibirica Ledeb. var. semenovii (B. Fedt-
of the range of the species. There are pure forests, sch.) T. S. Liu, Monogr. Gen. Abies: 188. 1971. Type:
but more often it is mixed with other conifers, e.g. Kirgyzstan: Talasskij Ala Tau, [“Turkestania, Bisch
Picea obovata, Larix gmelinii, in the mountains also Tasch”], V. A. Kallaur s.n., Sep 1897 (holotype G).
L. sibirica and Pinus sibirica; common broad-leaved
trees or shrubs are Betula pendula, Populus ­tremula,
Description
Sorbus aucuparia and Viburnum opulus. In the
120 southwestern part of its range other broad-leaved Shoots prominently ridged and grooved, yellowish
trees are mixed in: Tilia cordata, Ulmus scabra, and brown; buds slightly resinous. Resin ducts in leaves
Acer platanoides. small, marginal. Seed cones yellowish brown when
full grown but not dry.
Uses
Distribution
Siberian fir is an economically important timber
tree. Its wood is used in light-frame construction Kirgyzstan (Talasskij Alatau).
and for pulpwood. Planted in regions with mild TDWG codes: 32 KGZ
winters it can be damaged by ‘late’ frost; it is also in-
tolerant of air pollution. In Central and E Europe it
Ecology
has been introduced as an amenity tree and several
‘forms’ and cultivars are known. On N-facing slopes or in steep ravines at altitudes
between 1300 m and 2850 m a.s.l. where moisture
Two subspecies are recognized: from snowmelt remains available through summer.

Abies sibirica Ledeb. subsp. sibirica. Type: Russia:

Conservation
Altai Mts., C. F. von Ledebour s.n., 1826 (holotype
LE). IUCN: VU (D2)

Description
Abies spectabilis (D. Don) Mirb., Mém. Mus. Hist.
Shoots smooth or only faintly ridged, yellowish grey Nat. 13: 70. 1825. Pinus spectabilis D. Don, Prodr.
to fawn brown; buds resinous or very resinous. Res- Fl. Nepal. 2: 55. 1825. Type: India: Uttar Pradesh,
in ducts in leaves large, medial. Seed cones violet- Kumaun Himal, W. S. Webb ex herb. Wallich 6058
blue, maturing to bluish brown. (lectotype K, designated here). Fig. 18

Abies webbiana (Wall. ex D. Don) Lindl. var. brevi-

Distribution
folia A. Henry, in Elwes & Henry, Trees Gr. Brit. Ire-
Across N Russia, from Archangel’sk eastward to the land 4: 751. 1909; Abies spectabilis (D. Don) Spach
Amur River, southward to the mountains along the var. brevifolia (A. Henry) Rehd., J. Arnold Arbor. 1:
Sino-Russian border. 54. 1919.
TDWG codes: 14 RUE RUN 30 ALT BRY CTA IRK Abies spectabilis (D. Don) Spach var. langtangensis
KRA TVA WSB YAK 31 AMU KHA 36 CHX Silba, Phytologia 68: 22. 1990.

Conservation Etymology
IUCN: LC The species epithet means ‘splendid or ‘superb’.
 Vernacular names Distribution
Webb fir, Himalayan fir; bang, chilrao (Hindi); ba- Afghanistan: Hindu Kush; Pakistan: Karakoram
dar, paludar (Kashmir); dhunsing (Bhutan, Sikkim) Range; China: W Xizang [Tibet]; India: Kashmir
Himalaya; Nepal.
TDWG codes: 34 AFG 36 CHT 40 NEP PAK WHM-
Description
HP WHM-JK WHM-UT
Trees to 40–50 m tall, d.b.h. to 1.5–2.5 m; trunk
monopodial, straight, columnar, terete, stunted at
Ecology
tree line; crown broad pyramidal or columnar, flat
topped in old trees. Bark of young trees smooth, Abies spectabilis grows in the high mountains of 121
light grey or pinkish grey, later rough, breaking into the Himalayas, between 1600 m and 4000 m a.s.l.,
plates and furrowed in old trees. Branches of first or- but commonly in a ‘cloud belt’ between 2600 m
der thick, long, ascending near the top, curved down and 3800 m on N or NE slopes. The soils are vari-
below; branches of second order spreading, assur- ous alpine lithosols with good drainage. The climate
gent. Leading shoots thick, side shoots (branchlets) is cool, moist monsoon, with abundant precipita-
slender, firm, light yellowish brown, turning grey- tion, much of it as snow, in some parts of the west-
brown; surface prominently ridged and grooved, ern range rather drier. It forms pure stands at the
pubescent in grooves with brown hairs, but leading higher elevations up to the tree line, but usually it
shoots sometimes glabrous; leaf scars circular ovate. is mixed with conifers, e.g. Picea smithiana, Pinus
Vegetative buds ovoid or subglobose, large, on lead- wallichiana, Juniperus wallichiana, and in Nepal also
ing shoots 10 × 8 mm, densely covered with resin; with Tsuga dumosa. It is usually well separated in
bud scales triangular, keeled, with erose margins, elevation from the lower occurring fir A. pindrow.
brown or red-brown, persisting several years. Leaves Juniperus squamata (a shrub), Rhododendron cam-
spirally arranged, pectinate in two lateral sets, the panulatum and Betula utilis exceed A. spectabilis
shorter upper leaves directed forward but leav- in elevation at tree line; at lower elevations some
ing shoot bare, on coning shoots assurgent, 2.5–6 broad-leaved trees appear, e.g. Juglans regia, Quercus
cm long, 2.2–3.5 mm wide, twisted at base, linear, spp., Acer spp., Prunus spp., Ulmus spp., and Aescu-
flattened, with slightly recurved margins, continu- lus indica, and in Nepal especially large species of
ously grooved above, dark glossy green above, white Rhododendron.
bands below, midrib and margins glaucous green;
apex emarginate or bifid, on coning shoots obtuse.
Conservation
Stomata in two broad bands separated by a midrib
below. Pollen cones lateral, crowded, pendulous, 3–4 IUCN: LC
cm long, yellowish, with purplish blue microsporo-
phylls. Seed cones lateral, erect, nearly sessile, broad
Uses
cylindrical, with truncate apex, (8–)10–17 cm long,
4–7 cm wide, violet-blue when immature, maturing Webb fir is an important timber tree in the Hima-
to dark grey-blue, ripening to dark purplish brown; layas, where it is used in construction (house build-
cone rachis persistent, narrowly conical, dark bluish ing), in particular for interior work such as floor
brown. Seed scales flabellate-cuneate, thin, length boards, ceilings and stairs. In some parts shingles
× width at mid-cone 1.5–2.5 × 3–3.5 cm; surface are used for roofing; another use is the manufacture
smooth, slightly striated, puberulent on exposed of boxes to pack tea. Its uses are similar to those of
parts; upper margin entire, undulate or incurved; Pindrow fir, but as it occurs at higher elevations and
base pedicellate, laterally auriculate. Bracts spathu- often up to the tree line it is commercially of less
late, with rounded, serrulate margin and small cusp, importance. A local use of the purple cone scales has
1.5–2 cm long, usually included, or slightly exserted been to make a dye. Webb fir was introduced in Eng-
near base of cone. Seeds cuneate, 10 × 5 mm, brown; land in 1822 and remains in cultivation. The variety
seed wings cuneate-oblong, 12 × 5 mm, brown-­ brevifolia, as occasionally encountered in gardens
violet. and parks, is merely a high altitude form, as is the
type specimen of the species. In nature, the length
 of the leaves varies, not only with the altitude, but violet-brown; cone rachis persistent, fusiform, dark
also on the trees themselves. Describing and naming violet-brown. Seed scales cuneate-flabellate, length
taxa from an odd tree in cultivation often leads to × width at mid-cone 1.3 × 1.5 cm; apically thick-
superfluous names. ened; surface smooth, puberulent; upper margin
entire, incurved; base auriculate-pedicellate. Bracts
oblong-spathulate, 1.5–1.8 cm long, slightly exserted,
Abies squamata Mast., Gard. Chron., ser. 3, 39: 299, cusps recurved or straight. Seeds oblong-cuneate,
f. 121. 1906. Type: China: Sichuan, Daxue Shan, 5 × 2.5 mm, blackish brown; seed wings cuneate-
Kangding, [“W of Tachien-lu”], E. H. Wilson 3019 obovate, 7–8 × 6 mm, violet-brown.
(holotype K). Fig. 19, 20
122
Distribution
Etymology
China: S Gansu, S Qinghai (Baima Xian), W Si-
The species epithet refers to the scaly bark. chuan, E Xizang [Tibet] (Markam Xian).
TDWG codes: 36 CHC-SC CHN-GS CHQ CHT
Vernacular names
Ecology
Flaky fir; linpi leng shan (Chinese)
A subalpine species of the high mountains of west-
ern China, where it occurs between 3500 m and
Description
4500 m a.s.l. [3000–4700 m according to Liu (1971)]
Trees to 40 m tall, d.b.h. to 2 m; trunk monopodi- making it one of the highest reaching mountain
al, straight, columnar, terete; crown broad or nar- trees in the world. The soils are commonly grey-
row, conical. Bark of young trees smooth, purplish brown mountain podzols or lithosols. The climate
or pink brown, exfoliating like Betula, the bark in is cold, relatively dry (arid in E Xizang), but usually
older trees remaining shaggy from flakes of papery perpetual snow at higher elevations provides suffi-
bark, breaking into rough and hard plates, orange- cient moisture throughout the year. It is a constitu-
brown (when freshly exposed), blackish grey at base ent of mixed coniferous high altitude forests, with
of trunk. Branches of first order long, spreading among other species Abies recurvata, A. fargesii var.
horizontally, the lower pendant; branches of second faxoniana, Picea likiangensis var. rubescens, P. as-
order spreading horizontally, assurgent near the top perata, P. linzhiensis (in E Xizang), Larix potaninii
of the tree. Branchlets stout, firm, dark reddish or and possibly also Tsuga forrestii. There are very few
purplish brown, shining, ridged and grooved, usu- broad-leaved trees at these high elevations, Betula
ally glabrous or a few hairs in the grooves; leaf scars albosinensis and B. utilis var. prattii being the most
circular. Vegetative buds ovoid-globular, thickly common.
covered with white, soon eroding resin; bud scales
triangular-ovate, keeled, reddish brown, persisting
Conservation
several years. Leaves spirally arranged, dense, rigid,
spreading in two lateral, pectinate sets, assurgent At these high altitudes forests form isolated patch-
above, especially on cone bearing shoots, (1–)1.5– es on favourable sites, surrounded by treeless sub-
2.5(–2.8) cm long, 1.5–2 mm wide, curved or twisted alpine vegetation. Direct exploitation of the timber
at base, linear, more or less falcate, flattened, grooved in these forest remnants is easily unsustainable due
above, green or glaucous green above, two glaucous to very slow growth and has led to a decline of this
white or greenish white bands below; apex obtuse and other conifer tree species in these forests.
or acute. Stomata few to many near apex above, IUCN: VU (A1d)
in two bands separated by a midrib below. Pollen
cones lateral, on the underside of shoots, pendant,
Uses
2–3 cm long, yellowish, with purple microsporo-
phylls. Seed cones lateral, erect, short pedunculate, Flaky fir is a potential timber tree but its occurrence
ovoid-cylindric, with obtuse apex, 5–7 cm long, 3–4 at extremely high altitudes in inaccessible places
cm wide, violet-blue when immature, ripening to prevents it from being exploited commercially.
­Ernest Wilson collected this fir with its peculiar bark microsporophylls. Seed cones lateral, erect, often
in June 1904 in the Daxue Shan of western Sichuan, crowded, short pedunculate or almost sessile, cylin-
China, when on a plant hunting expedition for drical to ellipsoid (often irregular), with obtuse or
Veitch & Sons in England. Although it was sucess- papilliform apex, (3–)4.5–7.5(–8) cm long, (1.5–)2–
fully introduced in Europe and North America, it 2.5(–3) cm wide, dark bluish purple (rarely green or
has remained rare in cultivation, restricted to a few olive-green) when immature, ripening to blackish
collections in botanic gardens and arboreta, where brown or brown; cone rachis persistent, narrowly
it tends to be a slow grower. Its unusual bark has an conical, purplish brown. Seed scales narrowly reni-
attraction to dendrologists, but unless renewed seed form or almost crescent shaped, length × width at
collecting from wild sources can be resumed, this mid-cone 0.8–1 × 1.4–1.6 cm; surface smooth, (sil-
species may gradually disappear from horticulture. very) puberulent on exposed parts; upper margin 123
entire, incurved; base pedicellate. Bracts obcordate,
1–1.2 cm long, slightly exserted or only the cusps
Abies veitchii Lindl., Gard. Chron. 1861: 23. 1861. exserted, straight or recurved. Seeds cuneate, 5–6 ×
3 mm, greyish black tinged with green; seed wings
broadly cuneate, 3 × 5 mm, colour as seeds or black-
Etymology
ish purple.
This species has been named after John Gould
Veitch, founder of Veitch Nurseries, who ‘discov-
Distribution
ered’ it in 1861 on Mt. Fuji.
Japan: Honshu, Shikoku.
TDWG codes: 38 JAP-HN JAP-SH
Vernacular names
Veitch’s fir; shirabiso, shiro-momi (Japanese)
Ecology
The ‘typical’ variety of this species grows on high
Description
mountains at elevations between 1200 m and 2800
Trees to 25–30 m tall, d.b.h. to 0.8–1 m; trunk mo- m a.s.l. [reported from as low as 1050 m (Wilson,
nopodial, straight, columnar, terete; crown (nar- 1916)]. The soils are usually of volcanic origin, pod-
rowly) pyramidal. Bark of young trees smooth, with zolic and well drained. The climate is cool and wet,
prominent resin blisters, (light) greenish grey, lower with annual precipitation between 1000 mm and
part of trunk becoming scaly and dark grey. Branch- 2500 mm, and with cold, snowy winters; frequent
es of first order slender, relatively short, spread- typhoons cause destruction of the forest in most
ing horizontally, ascending near the top; branches places before it reaches an age of 250 to 300 years
of second order similar. Branchlets slender, firm, (Franklin et al., 1979). This variety is usually mixed
young shoots green or light brown, yellowish grey with other conifers, e.g. Abies mariesii, Picea jezoen-
in second year; surface smooth, in later years with sis subsp. hondoensis, Larix kaempferi, Thuja stand-
shallow grooves, densely yellowish pubescent, soon ishii, Pinus parviflora, at the highest elevations Pinus
glabrous; leaf scars circular, yellowish brown. Veg- pumila, and the ubiquitous Tsuga diversifolia. The
etative buds ovoid globular, 3 × 3 mm, resinous; most common broad-leaved trees are Betula erma-
bud scales triangular, with erose margins, reddish nii, Sorbus commixta, Prunus nipponica, and Acer
or purplish brown. Leaves spirally arranged, radi- spp. at lower elevations, and Betula corylifolia near
ally spreading, the lower leaves more or less pecti- the tree limit.
nate, the upper leaves directed forward, covering
shoot, on coning shoots slightly assurgent, (0.5–)
Uses
1.5–3(–3.8) cm long, 1.5–2.2 mm wide, widest near
the emarginate or truncate apex, linear-ligulate or Veitch’s fir is a relatively small tree which yields tim-
falcate, grooved above, flattened, dark green above, ber of low grade, mainly used for the manufacture
two whitish bands below. Stomata in two bands of paper pulp. It is fairly common in cultivation, as
separated by a midrib below. Pollen cones lateral, an amenity tree in parks and gardens and in collec-
axillary, pendant, 1–1.5 cm long, yellowish, with red tions (arboreta). Its popularity in the British Isles,
 where it was introduced in 1879 from seeds sent by
Conservation
C. Maries to Veitch & Son in Chelsea, England, was
due to the suitable and similar climate. As happens IUCN: NT
in such cases, this popularity has waned as other
trees became available.
Abies vejarii Martínez, Anales Inst. Biol. Univ. Nac.
2 varieties are recognized: México 13 (2): 629. 1942.

Abies veitchii Lindl. var. veitchii. Type not

Etymology
designated.
124 Named after Octavio Véjar Vázquez, at the time
Abies veitchii Lindl. var. olivacea Shiras., Bot. Mag. Mexican Minister for Public Education.
(Tokyo) 27: 132. 1913; Abies veitchii Lindl. f. oliva-
cea (Shiras.) Cinovskis, Introd. Rast. Bot. Sad. Pri­
Vernacular names
baltiki: 33. 1974.
Abies veitchii Lindl. var. komagatakensis Hayashi, Vejar’s fir; hayarin (Mexico)
Bull. Govt. Forest. Exp. Stat. 57: 151. 1952.
Description
Description
Trees to 35–40 m tall, d.b.h. to 1–1.5 m; trunk mo-
Leaves 1.5–3(–3.8) cm long, 1.5–2.2 mm wide. Seed nopodial, straight, columnar, terete; crown broad
cones cylindrical, (4–)4.5–7.5(–8) cm long, 2–2.5 cm conical or pyramidal, old trees rather open. Bark of
wide; bracts slightly exserted beyond the cusps. young trees smooth, thin, grey, of old trees rough,
scaly and fissured on lower part of trunk, grey-
brown. Branches of first order long, slender, spread-
Distribution
ing horizontally, ascending near the top; branches of
Japan: Honshu, Shikoku. second order similar. Branchlets slender, firm, pur-
TDWG codes: 38 JAP-HN JAP-SH plish red in the first year, becoming lighter reddish
brown or orange-brown; surface almost smooth,
shallowly grooved, glabrous; leaf scars circular. Veg-
Conservation
etative buds globose, 3 × 2.5 mm, very resinous; bud
IUCN: LC scales triangular ovate, keeled, light brown, persist-
ing several years. Leaves spirally arranged, the lower
Abies veitchii Lindl. var. sikokiana (Nakai) Kusaka, ones pectinate, the other leaves directed forward,
Conif. Japon. Ill., ed. 2: 212. 1954. Abies sikokiana covering shoot, on coning shoots spreading radially,
Nakai, Bot. Mag. (Tokyo) 42: 452. 1928. Type not 1–2(–2.5) cm long, 1.3–2 mm wide, slightly twisted
designated. or curved at base, narrowly lanceolate-linear, shal-
lowly grooved above, flattened, grey-green or dark
green above, whitish with a green midrib and mar-
Description
gins below; apex acute or acutish. Stomata in a few
Leaves (0.5–)0.8–2 cm long, 2 mm wide. Seed cones or up to 10 lines above, in two bands separated by a
ellipsoid-cylindrical, 3–4 cm long, 1.5–2 mm wide; midrib below. Pollen cones lateral, crowded, nearly
only the bract cusps are exserted. globular, 0.5 cm, with reddish microsporophylls.
Seed cones lateral, erect, short pedunculate, barrel-
shaped or ovoid-oblong, with obtuse or truncate
Distribution
apex, 6–12(–15) cm long, 4–6(–7?) cm wide, dark
Japan: Shikoku. purple when immature, maturing to purplish or
TDWG codes: 38 JAP-SH bluish tinged with brown, becoming dark brown
when ripe; cone rachis persistent, cylindric-coni-
cal, brown. Seed scales cuneate-flabellate, length ×
width at mid-cone 1.5–2 × 2–2.5 cm; surface smooth,
puberulent on exposed parts; upper margin en- their importance as timber trees is negligible. The
tire, incurved; base pedicellate. Bracts oblanceo- species (including its infraspecific taxa) has been
late-spathulate, with a broad, triangular cusp with introduced to cultivation in the USA and Europe,
denticulate margins, 1.7–2.5 cm long, included or ex- but it remains restricted to arboreta and other plant
serted, if exserted at first erect, but recurved in ripe collections despite its attractiveness and suitability
cones. Seeds cuneate-oblong, 8–10 mm long, brown; especially in regions with warm summers and mild,
seed wings cuneate-triangular, 15 × 12 mm, brown wet winters. Cultivation from wild origin seed of var.
with a violet tinge. mexicana would under appropriate circumstances
contribute to ex situ conservation of this threatened
taxon.
Taxonomic notes
125
In a recent paper Strandby et al. (2009) carried out a 3 varieties are recognized:
morphometric study of Mexican and Central Amer-
ican firs, in which they concluded that Abies vejarii Abies vejarii Martínez var. vejarii. Type: Mexico:
cannot be retained as a separate taxon from A. religi- Tamaulipas, 20 km N of Miquihuana, M. Martínez
osa and should therefore be merged. They proposed 3531 (holotype MEXU).
the new combination A. religiosa subsp. mexicana to
accommodate populations formerly named A. ve-
Description
jarii, A. vejarii subsp. mexicana, A. guatemalen-
sis, A. tacanensis, A. zapotekensis and varieties of Seed cones 6–12 cm long, 4–6 cm wide; bracts ex-
A. guatemalensis. They admit that “a consensus on serted.
the taxonomy of Abies ocurring in southern Mexi-
co and Guatemala is undoubtedly still remote” and
Distribution
therefore this new taxonomy, reducing several spe-
cies to synonymy or infraspecific status, is merely NE Mexico: Sierra Madre Oriental in Coahuila,
noted here. Nuevo León and Tamaulipas.
TDWG codes: 79 MXE-CO MXE-NL MXE-TA
Distribution
Conservation
NE Mexico: Sierra Madre Oriental in Coahuila,
Nuevo León, Tamaulipas. IUCN: LC
TDWG codes: 79 MXE-CO MXE-NL MXE-TA
Abies vejarii Martínez var. macrocarpa Martínez,
Anales Inst. Biol. Univ. Nac. México 19: 90. 1948.
Ecology
Type: Mexico: Coahuila, Saltillo, Mesa de las
Abies vejarii is a high mountain species, occurring Tablas, E. E. M. Loock 123 (holotype MEXU).
between (2000–)2800 m and 3300 m a.s.l. on steep
mountain slopes near the summits or in cool ra-
Description
vines. The soils are usually poor in humus content,
but moist; the climate is cool, with relatively dry Seed cones 10–15 cm long, 5(–7?) cm wide; bracts
summers and wet winters. The species is commonly exserted.
associated with various species of Pinus and with
Pseudotsuga menziesii var. glauca, also with Quer-
Distribution
cus spp.; Cupressus arizonica and Picea engelman-
nii subsp. mexicana (another endemic relict taxon) Mexico: Coahuila (Mesa de las Tablas), Nuevo León
have been reported with Abies vejarii var. mexicana. (Cerro Potosí).
TDWG codes: 79 MXE-CO MXE-NL
Uses
Conservation
This species and its infraspecific taxa are relatively
rare trees with limited distribution, consequently IUCN: LC
 Abies vejarii Martínez var. mexicana (Martínez)
Vernacular names
T. S. Liu, Monogr. Gen. Abies: 261. 1971. Abies
mexicana Martínez, Anales Inst. Biol. Univ. Nac. Yuanbaoshan fir; Yuanbaoshan lengshan (Chinese)
México 13 (2): 626. 1942; Abies vejarii Martínez
subsp. mexicana (Martínez) Farjon, Pinaceae
Description
(Regnum Veg. 121): 103. 1990. Type: Mexico: Nuevo
León, Sierra de Santa Catarina, M. Martínez s.n., Trees to 25 m tall, d.b.h. to 0.6 m (or more); trunk
1939 (holotype MEXU). monopodial, straight, columnar, terete; crown prob-
ably like A. forrestii (not described). Bark smooth
at first, becoming ridged and grooved, divided into
Description
126 small plates in old trees. Branches of first order long,
Seed cones to 10 cm long; bracts with a narrow, spreading horizontally; branches of second order
straight cusp, entirely included. spreading and ascending. Branchlets slender, firm,
yellowish brown or light brown, becoming light or
dark brown, glabrous; leaf scars circular. Vegetative
Taxonomic notes
buds ovoid-conical, very resinous; bud scales red-
Rushforth (1987) has rejected alliance of this variety dish brown, persisting several years. Leaves spirally
with A. vejarii and placed this taxon close to Abies arranged, densely set, subradially spreading above
durangensis. shoot, the lower leaves spreading laterally, the up-
per leaves shortest, on coning shoots assurgent or
erect, 1–2.7 cm long (leaves of young trees longer:
Distribution
3–3.8 cm, usually more pectinately arranged in two
Mexico: SE Coahuila, Nuevo León (Sierra Santa rows of equal length), longest in the middle portion
Catarina). of shoot, 1.8–2.5 mm wide, twisted or curved at base,
TDWG codes: 79 MXE-CO MXE-NL linear or ligulate-linear, flattened; margins (in sicco)
revolute, with a median groove above, dark green
above, two very white bands below; apex obtuse to
Ecology
slightly emarginate. Stomata absent above, in two
This variety occurs usually on N-facing slopes and broad bands separated by a midrib below. Pollen
in steep ravines at altitudes between 2000 m and cones solitary, axillary, on short branches, oblong
3000 m a.s.l. Cupressus arizonica and Picea engel- cylindrical, 1–1.5 cm long, yellow. Seed cones later-
mannii subsp. mexicana (another narrow endemic al, erect, short pedunculate to almost sessile, broad
relict taxon) have been reported to occur with A. ve- ovoid-cylindric, short, with slightly narrowed, ob-
jarii var. mexicana. tuse apex, 8–9 cm long, 4.5–5 cm wide, green or
yellowish green when immature, maturing to light
yellowish brown, becoming pale (yellowish) brown
Conservation
when ripe; cone rachis persistent, fusiform or coni-
IUCN: VU (D2) cal, brown. Seed scales cuneate-flabellate, length
× width at mid cone 2 × 2.2 cm; surface smooth,
with thickened central portion of apical part, ex-
Abies yuanbaoshanensis Y. J. Lu & L. K. Fu, Acta posed part densely greyish white puberulent; upper
Phytotax. Sin. 18 (2): 206. 1980. Type: China: margin entire, rounded to almost truncate, slightly
Guangxi, Rongshui Xian, Yuanbao Shan, Y. J. Lü incurved; lower margins auriculate; base short pedi-
1001 (holotype PE). cellate. Bracts large, broad oblong, widest in the
middle (9 m), with rounded, erose-denticulate up-
per margins and a small cusp, 2.5–2.8 cm long, of the
Etymology
same colour as the seed scales, apical part 6–7 mm
The epithet refers to the mountain where this spe- wide, reflexed, distinctly exserted. Seeds cuneate-
cies was discovered. oblong or obovate-oblong, 10 × 4 mm, resinous,
dark red brown; seed wings cuneate-dolabriform,
with a truncate apex, slightly longer than the seeds
Etymology
(9–11 × 8–10 mm), shining light brown.
The epithet refers to the name of the municipality
(Ziyuan Xian) in which the type specimen was col-
Taxonomic notes
lected.
This species is closely related to A. forrestii and is
therefore best placed in the section Pseudopicea,
Vernacular names
subsection Delavayianae.
Ziyuan fir; Ziyuan lengshan (Chinese)
Distribution
127
Description
China: N Guangxi (Rongshui Xian, Yuanbao Shan).
TDWG codes: 36 CHS-GX Trees to 30 m tall, d.b.h. to 0.6–0.9 m; trunk mono­
podial, straight, columnar, terete; crown broad,
conical or flat topped in old trees. Bark of young
Ecology
trees smooth, grey, in old trees shallowly ridged
The highest mountains in Guangxi, like Yuanbao and grooved, broken into small plates on lower part
Shan, have a very cool, wet climate, with annual of trunk. Branches of first order long, spreading
precipitation exceeding 2000 mm. The summers horizontally, ascending towards the top of the tree;
are cool and cloudy, the winters last 4–5 months branches of second order spreading or ascending.
and bring abundant snow from December through Branchlets thick, stout, at first light yellowish brown,
March. This species occurs in mixed deciduous-­ in the third year greyish brown; surface ridged and
coniferous forest with other conifers (e.g. Tsuga grooved between the leaves, glabrous or with short
chinensis) and broad-leaved trees dominated by hairs in the grooves; leaf scars circular. Vegetative
members of the Fagaceae; the Abies trees are very buds cylindric or ovoid-oblong, with acutish apex,
scattered. covered with a thin layer of white resin; bud scales
triangular, dorsally keeled, light yellowish brown.
Leaves spirally arranged, spreading laterally in two
Conservation
overlapping sets, of unequal length, the longest near
This species has an extremely limited distribution, base of shoot, on coning shoots upper leaves assur-
distant from other species, and is only known from gent, all leaves on vegetative shoots of young trees
one small area in Guangxi Province. pectinate, widely spaced, 2–4.8 cm long, 3–3.5 mm
IUCN: CR (B1+2a) wide, twisted or curved at base, linear, straight or
curved, flattened, with margins (in sicco) slightly re-
curved, dark green above, two greenish white bands
Uses
below; apex obtuse or slightly emarginate. Stomata
No uses have been recorded of this species. absent on the adaxial (upper) surface, in two bands
divided by a midrib below. Pollen cones lateral, in
leaf axils, ca. 2 cm long, yellowish, with red micro-
Abies ziyuanensis L. K. Fu & S. L. Mo, Acta sporophylls. Seed cones lateral, with 0.5–1 cm long
Phytotax. Sin. 18 (2): 208. 1980. Abies fabri (Mast.) peduncles, oblong-cylindric or elliptical, with ob-
Craib var. ziyuanensis (L. K. Fu & S. L. Mo) Silba, tuse apex, 10–11 cm long, 4.2–4.5 cm wide, green-
Phytologia 68: 14. 1990; Abies beshanzuensis ish or yellowish green when immature, maturing to
M. H. Wu var. ziyuanensis (L. K. Fu & S. L. Mo) dark greenish brown, becoming dark brown when
L. K. Fu & Nan Li, Novon 7 (3): 261. 1997. Type: ripe; cone rachis persistent, narrowly conical. Seed
China: Guangxi, Ziyuan Xian, Yinzhulao Shan, scales broad cuneate-flabellate, length × width at
Y. J. Lü 78001 (holotype PE). mid cone 2.3–2.5 × 3–3.3 cm; surface smooth, dark
brown when ripe, sparingly puberulent; upper mar-
Abies dayuanensis Q. X. Liu, Bull. Bot. Res. North- gin entire, rounded, not incurved, light brownish
East. Forest. Inst. 8 (3): 85. 1988. green; lateral margins finely toothed, auriculate near
the pedicellate base. Bracts oblong-spathulate, near
 apex 9–10 mm wide, near base 3 mm, 2.1–2.3 cm TDWG codes: 36 CHS-GX CHS-HN
long including the small cusp, included, or slight-
ly exserted near base of cone and recurved; apical
Ecology
margin with fine teeth. Seeds cuneate-oblong, ca. 10
× 4 mm, purplish grey with dark resin; seed wings Abies ziyuanensis is a rare fir occurring on the high-
cuneate-dolabriform, with rounded or slightly trun- est mountains in Guangxi and on the border with
cate apex, 13–15 × 12–14 mm, light purplish grey, Hunan, in a narrow belt between 1650 m and 1750
with dark spots, lustrous. m a.s.l. These mountains have a cool, very wet cli-
mate, with a mean annual temperature between
9.2–12 °C, and a winter period of 4–5 months (No-
Taxonomic notes
128 vember–March) in which the mean temperature is
According to Fu, Lu & Mo (1980), this species is between –3° to –5 °C (min. –10°). The weather is
closely related to A. beshanzuensis M. H. Wu, an- usually cloudy, with much fog, the annual precipi-
other novelty described around that time. Farjon & tation is 2100–2400 mm and snow lasts from De-
Rushforth (1989) classified A. ziyuanensis in section cember through March. Abies ziyuanensis occurs,
Momi, subsection Holophyllae, as its close affinity to together with other conifers, scattered in a mixed
A. chensiensis seemed more likely on morphological forest dominated by deciduous broad-leaved trees.
grounds. In Flora of China 4: 50 (1999) A. ziyuanen- Above 1700 m on Yuanbao Shan it is replaced by
sis has been reduced to a variety of A. beshanzuensis. A. yuanbaoshanensis.
If this taxonomy is accepted, A. beshanzuensis would
no longer be considered an extremely rare species,
Conservation
but merely a variety of a still rare, but more wide-
spread species. A recent phylogenetic study (Xiang This species is only known from less than five locali-
et al., 2009) based on nuclear DNA sequence data ties, some in close proximity. As it occurs lower on
(ITS regions) sampled A. ziyuanensis amongst 48 the mountains, it is potentially in danger of being
species. This species appeared to be closely related logged for local use of timber.
to A. homolepis and also to A. chensiensis. IUCN: CR (B1+2a)

Distribution Uses
China: NE Guangxi (Rongshui Xian, Yuanbao No uses have been recorded of this fir.
Shan), SW Hunan (Ziyuan Xian, Xingni, Chenbu).
Acmopyle Pilg., in Engler, Pflanzenreich 4 (5, 18): 117. 1903. Type: Acmopyle pancheri
(Brongn. & Gris) Pilg. (Podocarpaceae).

Greek: akme = highest point, tip; pyle = opening Acmopyle pancheri (Brongn. & Gris) Pilg., in
(gate); referring to the erect position of the mature Engler, Pflanzenr. IV.5 [18]: 117. 1903. Dacrydium
seed. pancheri Brongn. & Gris, Bull. Soc. Bot. France
16: 330. 1869; Nageia pancheri (Brongn. & Gris)
Kuntze, Revis. Gen. Pl. 2: 800. 1891. Type:
Description
New Caledonia: Grande Terre, Province Sud,
Dioecious (or sometimes monoecious?), small ever­ J. A. I. Pancher s.n. (holotype P). Fig. 21 129
green trees. Branching sparse in remote pseudo-
whorls (Massart’s model). Terminal buds absent. Podocarpus pectinatus Pancher ex Brongn. & Gris,
Leaves spirally arranged, of two kinds: small and Bull. Soc. Bot. France 16: 330. 1869.
scale-like on leading and fertile shoots, larger and Acmopyle alba J. T. Buchholz, Bull. Mus. Hist. Nat.
foliate on lateral, vegetative shoots; the larger leaves (Paris), sér. 2, 21: 281. 1949.
bilaterally flattened, falcate-linear, spreading ob-
liquely or in pinnate ranks on shaded shoots, single-­
Etymology
veined. Pollen cones lateral or terminal, axillary,
solitary or a few together, catkin-like; microsporo- This species was named after J. A. I. Pancher, who
phylls helically attached to a slender rachis on very collected the type specimen in 1869.
short stalks, triangular, with two basal pollen sacs
containing bisaccate pollen. Seed cones solitary or
Vernacular names
occasionally grouped, on scale-leaved pedunculate
branchlets axillary or sub-terminal on leafy shoots, No common names are recorded for this species.
when mature forming an irregular, fleshy and ver-
rucose receptacle from several unfertilized bracts
Description
which remain partly visible. Seeds single, from a
subterminal, inverted ovule, becoming (nearly) Small to medium size trees to 25 m tall, usually
erect at maturity, entirely enclosed by a fleshy epi- smaller; trunk monopodial, to 50 cm d.b.h. Bark on
matium, its base partly enclosed by the receptacle, trunk becoming hard and scaly, breaking into small
bluish pruinose when mature. plates, brown weathering grey; inner bark more
or less fibrous. Branches spreading horizontally in
2 species. young trees, irregular and assurgent in older trees,
with foliage towards the ends. Foliage branches
terete, with small spreading leaf apices terminating
Distribution
spirally arranged, decurrent scale leaves; ultimate
New Caledonia; Fiji (Viti Levu). foliage branchlets 3–15 cm long, alternating, mostly
plagiotropic, becoming deciduous after a few years,
with short scale leaves at the base which gradually
Key to the species of Acmopyle
enlarge to pectinately arranged foliage leaves. Foli-
– Leaves falcate to weakly S-curved, in middle age leaves on seedlings alternate, pectinate, linear,
of branchlets 10–30 mm long, 1.5–3 mm wide. straight, in the middle of the branchlet ca. 15 mm
Pollen cones elongating to 20–25 mm long, ca. long, 1.5 mm wide, acute or pungent. Foliage leaves
3 mm wide. Fully developed receptacles 15–20 × on young trees and mature trees similar, alternate,
8–10 mm. New Caledonia A. pancheri pectinate at 60–90°, straight or more often falcate
– Leaves straight or falcate to weakly S-curved, in towards apex, or on vigorous branchlets weakly S-
middle of branchlets 10–25 mm long, (0.6–)2– curved, in the middle of branchlets 10–30 mm long,
4(–4.8) mm wide. Pollen cones elongating to 1.5–3 mm wide, gradually shorter especially towards
5–8 mm long, ca. 1.5 mm wide. Fully developed the proximal and less so towards the distal end of
receptacles 7–9 × 7–8 mm. Fiji A. sahniana branchlets, twisted and decurrent at base; margins
 revolute; apex obtuse or acute, curved forward; mid-
Uses
rib narrow above, often faint, prominent below; leaf
colour lustrous dark green above, with two whitish No uses have been recorded of this species and it is
bands separated by a green midrib and with green in cultivation only in a few botanic gardens.
margins below. Stomata on both surfaces, in numer-
ous intermittent lines, some on the midrib on the
lower (abaxial) side and a few stomata near the base Acmopyle sahniana J. T. Buchholz & N. E. Gray,
and/or apex on the upperside. Pollen cones subter- J. Arnold Arbor. 28: 142. 1947. Type: Fiji: Western
minal or terminal on lateral foliage shoots, solitary Division, Viti Levu, Vakarogasiu Mountain, [“Na­
or 2–3 together, on short, scale-leaved peduncles, mosi Province”], J. W. Gillespie 3273 (holotype A).
130 initially globose, elongating to 20–25 mm long, ca.
3 mm wide at anthesis; microsporophylls imbricate,
Etymology
carinate, acute at first then acuminate, speading but
apically incurved, with two basal, small pollen sacs. This species was named after Prof. Birbal Sahni from
Seed cones subterminal or terminal on lateral foli- Lucknow, India, who studied the morphology of the
age shoots, sometimes axillary to scale leaves or foli- genus Acmopyle.
age leaves, solitary or sometimes 2–3 together, on up
to 12 mm long, curved, scale-leaved peduncles; com-
Vernacular names
posed of several sterile and 1–2 fertile, amalgamated
bracts, forming a fleshy, swollen, irregularly shaped, No common names have been recorded for this spe-
verrucose and pruinose receptacle 15–20 mm long cies.
and 8–10 mm wide. Seeds 1–2 at the distal end of
the receptacle, subglobose, 7–10 mm diam. includ-
Description
ing the covering epimatium, crested and pruinose
when full-grown. Small (monoecious?) trees to 12 m tall; trunk nor-
mally monopodial, to 20 cm d.b.h. Bark mostly
smooth, sometimes pustulate, brown weathering
Distribution
grey; inner bark red and more or less fibrous. Branch-
New Caledonia (Grande Terre). es sparse, spreading to form a crown half as wide as
TDWG codes: 60 NWC the tree height, with foliage towards the ends. Foli-
age branches terete, with very small spreading leaf
apices terminating spirally arranged, decurrent scale
Ecology
leaves; ultimate foliage branchlets 2–6(–12) cm long,
Acmopyle pancheri occurs scattered in rainforest as alternating, mostly plagiotropic, becoming decidu-
an understorey tree and in low forest and vegetation ous after a few years, with short scale leaves at base
bordering on ‘maquis minier’ as a small canopy tree. which often abruptly change to pectinately arranged
Its altitudinal range is from near sea level to at least foliage leaves. Foliage leaves on young trees and
1200 m a.s.l. and it is found on ultramafic soil de- mature trees similar, alternate, pectinate at 60–90°,
rived from serpentine or similar rock as well as on straight or more often falcate towards apex, or on
acidic soil from metamorphic schist. It is associated vigorous branchlets weakly S-curved, in the middle
with other conifers such as Araucaria spp., Dacry- of branchlets 10–25 mm long, (0.6–)2–4(–4.8) mm
dium araucarioides and Agathis ovata, as well as with wide, smallest towards proximal and distal end of
numerous angiosperms. branchlets, twisted and decurrent at base; margins
revolute; apex acute, curved forward; midrib nar-
row above, often faint, prominent below; leaf colour
Conservation
lustrous dark green above, with two whitish bands
IUCN: NT separated by a green midrib and with green mar-
gins below. Stomata on both surfaces, in numerous
i­ ntermittent lines, some on the midrib on the lower TDWG codes: 60 FIJ
(abaxial) side and a few stomata near the base and/
or apex on the upperside. Pollen cones subterminal
Ecology
or terminal on lateral foliage shoots, solitary or in
pairs, sessile or on very short peduncles, initially Acmopyle sahniana is a small tree occurring sparsely
globose, elongating to 5–8 mm long, ca. 1.5 mm in low rainforest on mountain ridges and summits
wide at anthesis; microsporophylls imbricate, cari- in a single locality on the island of Viti Levu. The
nate, acute, speading, with two basal, small pollen altitudinal range is from 600 m to 1050 m a.s.l.
sacs. Seed cones subterminal or terminal on lateral
foliage shoots, sometimes axillary to scale leaves or
Conservation
foliage leaves, solitary, on up to 6 mm long, curved, 131
scale-leaved peduncles; composed of 2–3 sterile and The very limited area of occupancy (AOO) of this
1(–2) fertile amalgamated bracts, forming a fleshy, species and an estimated population not exceeding
swollen, irregularly shaped, verrucose and green or 250 mature trees put this species in the Endangered
purple receptacle 7–9 mm long and 7–8 mm wide. category. Proposed mining activities in the area,
Seeds solitary at the distal end of the receptacle, which is not protected, further threaten this species
ovoid, 7–9 mm long, 5–6 mm wide including the (Doyle in Farjon & Page, 1999).
covering epimatium, striated and greyish violet with IUCN: EN (D)
whitish bloom when full-grown.
Uses
Distribution
No uses have been recorded of this species.
Fiji Islands (Viti Levu, Mt. Evans Forest Park).
 Actinostrobus Miq., in Lehmann, Pl. Preiss. 1: 644. 1845. Type: Actinostrobus
pyramidalis Miq. (Cupressaceae).

Greek: actino- = rayed, star-like; strobus = cone; re- 2a. Seed cone scales with straight, acute apex
ferring to the six scales coming together at the apex A. arenarius
of the seed cone. 2b. Seed cone scales with incurved, obtuse apex
A. pyramidalis
Description
132 Evergreen, monoecious (decumbent) shrubs or Actinostrobus acuminatus Parl., Index Sem. Hort.
trees with smooth, thin, flaking bark. Resin cavi- Florent. 1862: 25. 1862. Type: Australia: Western
ties in leaves. Branches short, stiff, spreading or Australia, [“between Moore & Murchison Rivers”;
ascending, forming a conical, pyramidal or bushy “Swan River”], J. Drummond 225 (holotype FI).
crown. Fastigiate forms common in nature. Leaves
in whorls of 3, decurrent; juvenile acicular leaves on
Etymology
young plants only or also on mature plants; adult
leaves mostly shorter than 5 mm, linear-lanceolate, The species epithet refers to the shape of the cone
abaxially keeled, denticulate on margins and keel apex (Latin: acuminatus = tapering from inwardly
or only on margins, acuminate-pungent, green or curved sides to a narrow point).
glaucous green, amphistomatic, stomata in 2 lines
on each face. Pollen cones small, cylindrical; micro-
Vernacular names
sporophylls 10–18, in whorls of 3, with 2–4 abaxial
pollen sacs. Seed cones solitary, with 2–3 together or Dwarf cypress
aggregated along branches and stems, when closed
broadly globose with dome-shaped apex, broadly
Description
globose-conical, or broadly utriculate. Bract-scale
complexes in two whorls of 3, of nearly equal size Shrubs, erect or decumbent to ascending, dense,
at maturity, subtended by 4–6 alternating whorls of 1–4.5 m tall; trunk short or multistemmed. Bark
3 broad, imbricate scale leaves. Bract tips entirely smooth, soon flaking, thin, brown-grey. Branches
included in cone scales. Cone scales oblong, open- numerous, spreading or prostrate, persistent, form-
ing valvately, with light coloured seed scars towards ing a broadly conical or more irregular crown (of-
base. Columella a strong, acute spike. Ovules in 2 ten partly buried in blown sand). Foliage branches
whorls of 4–6 axillary to bracts, erect. Seeds 8–12 per numerous, contorted, spreading irregularly or as-
cone, with 3 wings. Seedlings with 2 cotyledons. cending, rigid, slender, short, rather sturdy, angular
with decurrent leaf bases, persistent, grey-brown to
3 species. grey when leaves weather away. Leaves in alternat-
ing whorls of 3, decurrent; juvenile leaves on young
and mature plants, acicular, 10–20 × 0.8–1.5 mm,
Distribution
with spreading free part (patent), abaxially keeled,
SW Western Australia, in a narrow coastal strip acute-pungent, light green or yellowish green, adult
from S of Shark Bay to near Albany. leaves linear-lanceolate, 2–4 × 1 mm, with recurved
or incurved free apex, abaxially keeled, denticulate
on margins, acuminate, green; stomata in 2 nar-
Key to the species of Actinostrobus
row lines on each face. Pollen cones terminal on
1a. Both juvenile, long acicular and adult, scale ultimate branchlets, yellowish green turning light
leaves present on mature plants. Seed cones not brown, 7–10 × 2–3 mm; microsporophylls 10–16, in
in clusters on stems A. acuminatus whorls of 3, ovate with acute apex, slightly keeled
1b. Leaves on mature plants short, scale-like. Seed abaxially, with 3–4 abaxial pollen sacs. Seed cones
cones in clusters on stems 2 terminal on short (ca. 1 cm), slightly thickened, lat-
eral, short-leaved branchlets, mostly solitary or with Actinostrobus arenarius C. A. Gardner, J. Roy.
2–3 together, when closed broadly utriculate, with Soc. W. Austral. 47: 54. 1964. Actinostrobus pyra­
a distinct ‘neck’ below apex of scales, 15–25 × 10–20 midalis Miq. var. arenarius (C. A. Gardner)
mm, glaucous green maturing to brown. Bract-scale Silba, Phytologia Mem. 7: 11. 1984; Actinostrobus
complexes whorled, 6, of nearly equal size at matu- pyramidalis Miq. subsp. arenarius (C. A. Gardner)
rity, subtended by 4–6 alternating whorls of 3 broad, Silba, J. Int. Conifer Preserv. Soc. 13 (1): 1. 2006.
imbricate scale leaves with denticulate margins; Type: Australia: Western Australia, Tammin,
apex free, acuminate; smaller lower whorls partly C. A. Gardner 610 (holotype PERTH). Fig. 22, 23
overlapping larger upper whorls. Bract tips entirely
included in cone scales. Cone scales oblong, 15–20
Etymology
× 5–10 mm, more or less concave and finely wrin- 133
kled, dark brown abaxially, with recurved, acute The species epithet refers to its habitat (Latin: arena
apex, with smooth adaxial face, blackish or purplish = sand).
black, with light coloured seed scars towards base.
Columella a strong, acute central spike. Seeds 8–12
Vernacular names
per cone, only a small number fertile, about half of
these smaller and situated between bases of cone Bruce cypress
scales, irregularly triangular/angular, 5–6 mm long
(including wings 9–12 mm), yellowish brown; wings
Description
narrow, 2–3 mm wide.
Erect shrubs or small trees, dense, 2–5 m tall; trunk
short or multistemmed. Bark smooth, soon flak-
Distribution
ing, thin, brown-grey. Branches numerous, spread-
SW Australia: from Three Springs south to Perth ing or ascending, persistent, forming a narrowly or
Region, in proximity of the coast. broadly conical or more irregular crown. Foliage
TDWG codes: 50 WAU-WA branches numerous, contorted, spreading irregular-
ly or ascending, rigid, slender, short, rather sturdy,
angular with decurrent leaf bases, persistent, grey-
Ecology
brown to grey when leaves weather away. Leaves in
Mostly in low dwarf scrub (‘kwongan’), with Myrta- alternating whorls of 3, decurrent (juvenile leaves
ceae, Proteaceae etc.; also in low woodland domi- on young plants only), linear-lanceolate, 3–5(–9) ×
nated by Eucalyptus spp. (‘mallee’), with Banksia 1–1.5(–2) mm, with spreading or recurved free distal
attenuata, Allocasuarina fraseriana, Hakea conchi­ part, abaxially keeled, denticulate on margins and
folia, and Calothamnus quadrifidus; on plains in dry keel, acuminate-pungent, green or glaucous green;
or moist leached white or grey sand or clay usually stomata in 2 lines on each face. Pollen cones termi-
over laterite (hardpan). The climate is characterized nal on ultimate branchlets, yellowish green turning
by warm, dry summers and winter rainfall. light brown, 5–7 × 2–2.5 mm; microsporophylls 12–
18, in whorls of 3, ovate with cuspidate apex, slightly
keeled abaxially, with 2–4 abaxial pollen sacs. Seed
Conservation
cones terminal on short (0.5–1 cm), slightly thick-
Although its habitat has been altered in some areas ened, lateral, short-leaved branchlets, mostly ag-
by agricultural development, much remains and this gregated along branches and stems, when closed
species is only at low risk of extinction at present. broadly globose-conical, 12–18 × 10–18 mm, glau-
An increase of the incidence of fires could be haz- cous green or pruinose, maturing to purplish brown
ardous if it occurred in key populations. or grey-brown. Bract-scale complexes whorled, 6,
IUCN: NT of nearly equal size at maturity, subtended by 4–6
alternating whorls of 3 broad, imbricate scale leaves
with denticulate margins; apex appressed, rounded,
Uses
mucronate; smaller lower whorls partly overlapping
No uses are recorded. larger upper whorls. Bract tips entirely included in
cone scales. Cone scales oblong, 10–15 × 5–8 mm,
 more or less flat, smooth, grey-brown abaxially, arable fields (‘wheat belt’) in disturbed vegetation;
with straight, acute apex, with smooth adaxial face, on plains in leached white or yellow (gravelly) sand,
purplish brown, with light coloured seed scars to- sandstone, loam or on laterite (hardpan); to a maxi-
wards base. Columella ca. 6 × 4 mm, triangular, mum altitude of 200 m a.s.l. The climate is charac-
brown. Seeds 8–12 per cone, only a small number terized by warm, dry summers and winter rainfall.
fertile, about half of these situated between bases of
cone scales, irregularly triangular/angular, 4–5 mm
Conservation
long (including wings ca. 10 mm), yellowish brown;
wings 2–3 mm wide. This species is still widespread in most of its natu-
ral range despite the fact that large areas have been
134 converted to agriculture. It is in fact colonising road
Taxonomic notes
verges and abandoned fields, often forming groves
This species is a segregate from A. pyramidalis, of of some extent much as some species of Juniperus
which the proposing author emphasised differences are known to do in the northern hemisphere. It is
in glaucousness of leaves and cones as well as the therefore considered not threatened.
shape of female cone scales. The leaves of A. are- IUCN: LC
narius can be green as well as glaucous green, result-
ing in more or less glaucous foliage or sometimes
Uses
green foliage that is indistinguishable from that
of A. pyramidalis. As in other conifers, glaucous- No uses are recorded; it may be planted locally as an
ness of foliage is highly variable and unreliable as ornamental. See for comments about its potentials
a taxonomic character. In A. arenarius the free under A. pyramidalis.
apical parts of leaves are nearly always spreading,
while in A. pyramidalis they are either spreading
or incurved, which gives the former species on the Actinostrobus pyramidalis Miq., in Lehmann,
whole a rougher appearance in its foliage. This is Pl. Preiss. 1: 644. 1845. Type: Australia: Western
certainly not a fully distinctive character. The shape Australia, Gordon River, [“et in depressis distric­
of the mature cone scales does distinguish this spe- tum Perth et Wellington”; the specimen in P has
cies from A. pyramidalis, and they are indeed larger “Riv. des Cygnes” = Swan River], L. Preiss 1311
when fully grown. These characters place the cone (holotype U).
as intermediate in shape and size between A. acumi-
natus and A. arenarius. The two species A. arenarius
Etymology
and A. pyramidalis appear to be largely allopatric
but occupy very similar habitats; A. acuminatus is The species epithet refers to the habit of this shrub.
mostly sympatric with A. arenarius (which has a
larger range) but the two species are usually found
Vernacular names
at different localities and usually occupy somewhat
different microhabitats. Swan River cypress, Swamp cypress

Distribution Description
SW Australia: from Lake Grace to the Murchison Erect shrubs or small trees, dense, 2–6 m tall; trunk
River. short or multistemmed. Bark smooth, soon flaking,
TDWG codes: 50 WAU-WA thin, brown-grey. Branches numerous, ascending or
nearly erect, persistent, forming a narrowly coni-
cal or pyramidal, less often rounded crown. Foliage
Ecology
branches numerous, contorted, spreading irregular-
Usually in low, open dwarf scrub (‘kwongan’)with ly or ascending, rigid, slender, short, rather sturdy,
Myrtaceae (e.g. Melaleuca), Proteaceae (e.g. Banksia, angular with decurrent leaf bases, persistent, grey-
Conospermum, Grevillea, Hakea), Acacia spp., Calli- brown to grey when leaves weather away. Leaves in
tris preissii, and C. roei; often on road verges through alternating whorls of 3, decurrent (juvenile leaves
on young plants only), linear-lanceolate, 2–5(–8) × or yellow sand, reddish lateritic soil (hardpan) and
1–1.5(–2) mm, with spreading or incurved free distal gravel. The climate is characterized by warm, dry
part, abaxially keeled, denticulate on margins and summers and winter rainfall.
keel, acuminate-pungent, green; stomata in 2 lines
on each face. Pollen cones subterminal on ultimate
Conservation
branchlets, reddish yellow turning reddish brown,
3–6 × 1.5–2 mm; microsporophylls 12–18, in whorls While still common, because it occupies the small-
of 3, orbicular-ovate with cuspidate apex, slightly est range of the three species of Actinostrobus, which
keeled abaxially, with 2–4 abaxial pollen sacs. Seed in part coincides with a major metropolitan area,
cones terminal on short (0.5–1 cm), slightly thick- A. pyramidalis is more at risk. Threats are urbanisa-
ened, lateral, short-leaved branchlets, mostly ag- tion, conversion of wildland to agriculture (limited) 135
gregated along branches and stems, when closed and especially increased incidence of fire. This spe-
broadly globose with dome-shaped apex, 12–15 × cies is therefore at low risk but Near Threatened as
12–16 mm, green or purplish green, maturing to it is likely that some decline of populations or their
purplish brown or grey-brown. Bract-scale com- size may have occurred. On the other hand it is also
plexes whorled, 6, of nearly equal size at maturity, capable of colonising new sites, but a comprehensive
subtended by 4–6 alternating whorls of 3 broad, im- assessment of the dynamics of these populations is
bricate scale leaves with denticulate margins; apex not available.
appressed, rounded, minutely mucronate; small- IUCN: NT
er lower whorls partly overlapping larger upper
whorls. Bract tips entirely included in cone scales.
Uses
Cone scales oblong, 10–14 × 5–6 mm, more or less
flat, smooth, grey-brown abaxially, with incurved, No uses are recorded; it may be planted locally as an
obtuse apex, with smooth adaxial face, purplish ornamental. The lack of interest in the horticultural
brown or nearly black, with lighter coloured seed use of this species and of similar-looking A. arenarius
scars towards base. Columella a 5–7 mm long, dark in their native country is remarkable; their habit as
brown to blackish spike. Seeds 8–12 per cone, only an often naturally columnar to pyramidal shrub and
a small number fertile, about half of these situated the attractive green or glaucous green foliage and
between bases of cone scales, irregularly triangular/ seed cones seem to give them much merit. I suspect
angular, 3–5 mm long (including wings 6–9 mm), that this is an omission with cultural backgrounds.
yellowish brown; wings ca. 2 mm wide. European (largely British) immigrants have created
English gardens with English design and plantings,
including manicured lawns that in the climate of
Distribution
much of Australia require constant and expensive
SW Australia: From Lynton District (Hutt River) watering. The native flora is considered ‘bush’ and
S to Albany District. while several of its more showy species, e.g. of the
TDWG codes: 50 WAU-WA Banksiaceae and Myrtaceae, have found their way to
gardens in Europe and beyond, Australian gardens
have to look like those at home, which at least sub-
Ecology
conciously is still England. Actinostrobus pyramida-
In low, open dwarf scrub (‘kwongan’), or in low lis should be able to do in Australian gardens what
woodland dominated by Eucalyptus spp. (‘mallee’); introduced ‘cypresses’ are doing there now, but with
often in road verges through farmland in disturbed far less input of pumped up water. It would also do
vegetation, associated with other Myrtaceae (e.g. well in regions with a Mediterranean type climate
Melaleuca), Proteaceae (e.g. Banksia, Conospermum, in California, Chile, southern Europe and South
Grevillea, Hakea), Acacia spp., Xanthorrhoea sp., ­Africa, taking care that it does not become invasive
Callitris preissii, and C. roei; on plains in dry soil, or in natural vegetation. To establish this, more de-
in or near (salt) marshes and lakes, on leached white tailed research into its ecology is desirable.
 Afrocarpus (J. T. Buchholz & N. E. Gray) C. N. Page, Notes Roy. Bot. Gard. Edin­
burgh 45 (2): 383. 1989. Podocarpus sect. Afrocarpus J. T. Buchholz & N. E. Gray,
J. Arnold Arbor. 29: 57. 1948. Type: Afrocarpus falcatus (Thunb.) C. N. Page [Taxus
falcata Thunb.] (Podo­carpaceae).

Latin: afro- = African; carpus = fruit; name given to met with universal recognition, in particular among
distinguish the genus from the related genus Podo- South African botanists. Some morphological dis-
carpus. tinctions stressed by Page, such as the lack of a succu-
136 lent and colourful receptacle, present in Podocarpus
sensu stricto, are in reality less rigidly distinct, with
Description
the situation found in P. henkelii in particular rep-
Evergreen, dioecious, large trees. Resin canals (1) in resenting an intermediate stage. The function of the
leaves only. Bark thin, becoming scaly with small receptacle, serving seed dispersal by means of offer-
plates. Terminal buds small or absent. Leaves spi- ing birds an attractive ‘imitation fruit’, is apparently
rally inserted or (on young plants) opposite, twisted taken over by the fleshy epimatium surrounding the
in opposite directions so as to orientate the blades seed in Afrocarpus. There are botanists who would
[on opposite sides of the same shoot] with the adax- be inclined to emphasize such adaptational traits as
ial side uppermost and the adaxial side downwards, significant, but others who would argue that they do
pectinately arranged or ascending, flattened, coria- not necessarily indicate phylogeny (adaptation can
ceous, narrowly lanceolate-elliptic to linear-lanceo- lead to convergent evolution) and if not, should be
late, with a single midrib. Stomata on both surfaces disregarded. Other morphological differences, such
(leaves amphistomatic). Pollen cones axillary, soli- as the amphistomatic leaves of the species in Afro-
tary or in groups of 2–3 on short naked peduncles, carpus, appear to be more consistent and have sup-
becoming narrowly cylindrical and catkin-like; ported a phylogeny separating Afrocarpus (Kelch,
micro­sporophylls spirally inserted, with two pollen 1997). Recently, research into this taxonomic ques-
sacs containing bisaccate pollen. Seed cones axil- tion has involved molecular (DNA) analysis (Barker
lary or just below foliage leaves on a short pedun- et al., 2004 and papers cited therein) and these data
cle with or without small scale leaves, consisting of also gave support to the recognition of Afrocarpus
a few sterile bracts and one larger fertile bract with as distinct from Podocarpus. Cladistic analyses of
an axillary, inverted ovule. Seeds single per reduced morphological as well as molecular data therefore
cone, subtended by small, withering scales (not by a appear to confirm the taxonomy proposed by Page,
receptacle), entirely enclosed by a fleshy, subglobose as is here adopted.
to obovoid or ellipsoid epimatium, maturing from
greenish to yellow or reddish brown; seed proper
Key to the species of Afrocarpus
with a hard, strongly sclerified seed coat.
The species of Afrocarpus are difficult to key out on
5 species. vegetative characters only; it is therefore necessary
to examine especially the seeds with and without
their soft covering (epimatium) and in their fully
Distribution
mature state in most cases.
E Africa: Ethiopia, Kenya, Uganda, Rwanda, Bu-
rundi, Congo Republic (Kivu), Tanzania, Malawi; 1a. Adult leaves 3–7(–8) mm wide, up to 11 cm long
S Africa: South Africa, Mozambique (Lourenço (usually to 8 cm long). Microsporophylls of
Marques), Swaziland; W Africa: São Thomé. pollen cones broadly triangular A. mannii
1b. Adult leaves (1.5–)2–4(–5) mm wide, up to 8 cm
long (usually to 5 cm long). Microsporophylls
Taxonomic notes
of pollen cones triangular-trullate 2
The genus Afrocarpus, raised to that rank by Page 2a. Buds very small, 0.6–1 mm diam. Midrib
(1989) from a section under Podocarpus, has not prominent on both sides of leaves. Seed coat
 (3–)4–6(–8) mm thick (remove epimatium) to a fine point. Adult leaves shorter, (2–)3–5(–6)
A. usambarensis cm long, (1.5–)2–4(–5) mm wide, spirally arranged,
2b. Buds larger, ca. 2 mm diam. Midrib prominent twisted at the attenuate base, spreading to ascend-
on the adaxial (upper) sides of leaves only. Seed ing, straight or rarely slightly falcate, linear-elliptic
coat 1–4 mm tick (remove epimatium) 3 or with parallel sides, gradually or abruptly tapered
3a. Seeds including the epimatium 20–30 mm above 3/4 of their length, with a raised midrib most
long; seed proper (remove epimatium) rugose, prominently on the adaxial (lower) side, grey-green;
with a 2–4 mm thick seed coat A. dawei apex acute. Stomata on both surfaces, arranged in
3b. Seeds including the epimatium 12–20(–23) mm numerous intermittent lines not well separated by
long; seed proper smooth or verrucose, with a the midrib. Pollen cones solitary or with 2–3 on very
1–1.5 mm thick seed coat 4 short stalks or subsessile, axillary to foliage leaves or 137
4a. Adult leaves (1–)2–4(–4.5) cm long. Seed prop- not, subtended by a whorl of papery bracts, initially
er spherical/compressed, 10–14 mm diam., with subglobose, elongating to cylindrical, 10–20(–25)
a verrucose seed coat A. falcatus mm long, 2.5–3.5 mm diam.; microsporophylls spi-
4b. Adult leaves (1.5–)3–6(–8) cm long. Seed prop- rally arranged, imbricate before anthesis, triangular-
er ovoid/compressed, 12–18 mm long, with a trullate, ca. 1 mm wide, with denticulate-lacerate
smooth seed coat A. gracilior margins and acute or apiculate apex, bearing two
subglobose pollen sacs. Seed cones solitary on small,
scaly or leafy branchlets situated axillary to or below
Afrocarpus dawei (Stapf) C. N. Page, Notes Roy. foliage leaves, with several sterile and one terminal,
Bot. Gard. Edinburgh 45: 384. 1989. Podocarpus larger fertile bract. Mature seed cones with a single
dawei Stapf, in Prain, Fl. Trop. Afrika 6 (2): 342. seed subtended by a single, short bract; with seed
1917. Type: Uganda: Nile Land, South Buddu, enclosed by a fleshy, firm epimatium that ripens
Kaigera River, M. T. Dawe 961 (holotype K) from green or glaucous green to yellow, globose or
[holotype K, not found; isotype B, destroyed]. subglobose, 25–30 mm long. Seed proper ovoid but
slightly compressed laterally, 16–21 mm long, 12–15
mm wide, with a rugose surface and hard, 2–4 mm
Etymology
thick seed coat.
The species epithet commemorates the botanist
Morley Thomas Dawe (1880–1943), who collected
Distribution
the type specimen.
Northern Tanzania (Kagera and Mara Prov.); Ugan-
da.
Vernacular names
TDWG codes: 25 TAN UGA
No vernacular names have been recorded for this
species.
Ecology
Afrocarpus dawei occurs as a co-dominant or
Description
emergent tree in seasonal swamp forest on or near
Trees to 33 m tall, d.b.h. to 1 m., with a long, clear bole. floodplains of slow running rivers E and S of Lake
Bark smooth in young trees, flaking in rectangular Victoria. It is most commonly associated with
or rounded small plates in large trees, dark brown, Baikiaea minor (Leguminosae) and Mimusops sp.
weathering grey. Branches ascending and spread- (­Sapotaceae). These flatland (elevation 1100–1200 m
ing, forming a small flat-topped crown. Foliage a.s.l.) river forests stand on recent river alluvial soils
dense, on numerous branches; new lateral branch- and are inundated in the rainy season, when they
lets ridged or more or less quadrangular; terminal become virtually inaccessible.
buds small, ca. 2 mm diam., or absent; bud scales
triangular or rounded with or without a rostrate
Conservation
apex. Leaves on seedlings and young plants mostly
opposite, narrowly linear-lanceolate, up to 17 cm This species does not fulfil the criteria for V, E, CR
long and 4–8 mm wide, straight or falcate, ­tapering based on GIS information; its extent of occurrence
 (EOO) is 97,974 km² (possible NT) and area of occu-
Description
pancy (AOO) is 18,215 km² (LC). Human Footprint
AVG in the area is low (7). It is considered NT on the Trees generally to 25 m tall but attaining 60 m,
basis of observational reports on exploitation cited with a massive trunk, d.b.h. to 2 m. Bark smooth in
on herbarium specimens, which indicates there is young trees, flaking in rectangular or rounded small
reason to suspect decline under criterion A2, but it plates in large trees, purplish brown or dark brown,
cannot be quantified. A notable population occurs weathering grey. Branches ascending and spreading
in the Minziro Forest Reserve in Tanzania, where 13 forming a broad, domed crown. Foliage dense, on
of 19 herbarium collections used in the assessment numerous branches; new lateral branchlets ridged
were made. or more or less quadrangular; terminal buds small,
138 IUCN: NT ca. 2 × 1 mm, or absent; bud scales narrowly triangu-
lar, acute. Leaves spirally arranged, on seedlings and
young plants narrowly linear-lanceolate, up to 12 cm
Uses
long and 3–6 mm wide, straight or falcate, tapering
The timber of this species, which can grow a tall, to a fine point. Adult leaves much shorter, (1–)2–4(–
straight bole without branches, is valued for con- 4.5) cm long, (1.2–)2–4(–5) mm wide, twisted at the
struction and carpentry or joinery work, and trees narrowed base, spreading to ascending, straight or
are singled out despite the seasonally difficult access slightly falcate, linear-lanceolate to linear-elliptic,
to be logged. It is not exported and used locally or with a conspicuously raised midrib adaxially (lower
regionally. The species is not known to be in cultiva- side), obscurely present abaxially, grey-green; apex
tion. acute to obtuse. Stomata on both surfaces, arranged
in numerous intermittent lines not well separated by
the midrib. Pollen cones solitary or with 2–4 on very
Afrocarpus falcatus (Thunb.) C. N. Page, Notes Roy. short stalks or subsessile, axillary to foliage leaves or
Bot. Gard. Edinburgh 45: 383. 1989. Taxus falcata not, subtended by a whorl of papery bracts, initial-
Thunb., Prodr. Pl. Cap.: 117. 1800; Podocarpus falca­ ly subglobose, elongating to cylindrical, 5–13(–15)
tus (Thunb.) Endl., Syn. Conif.: 219. 1847; Nageia mm long, 2–3 mm diam.; microsporophylls spirally
falcata (Thunb.) Kuntze, Revis. Gen. Pl. 2: 800. ­arranged, imbricate before anthesis, broadly trian-
1891 (nom. illeg. Art. 53.1); Decussocarpus falcatus gular-trullate, ca. 0.6 × 1 mm, with lacerate margins
(Thunb.) de Laub., J. Arnold Arbor. 50: 359. 1969. and acute or apiculate apex, bearing two slightly
Type: South Africa: Cape Province, [“e Cap. b. Sp. elongate pollen sacs. Seed cones solitary on small,
Ribeek(capel?), Vleermuysdrift”], C. P. Thunberg scaly branchlets situated axillary to or below foliage
UPS 23779 (holotype UPS). Fig. 24 leaves, with several sterile and one terminal, larger
fertile bract. Mature seed cones with a single seed
Podocarpus gracilimus Stapf, in Prain, Fl. Trop. Af- subtended by a single, short bract; with seed en-
rica 6 (2): 343. 1917. closed by a fleshy, firm epimatium that ripens from
glaucous green to yellow or light reddish brown,
Afrocarpus gaussenii (Woltz) C. N. Page, Notes Roy. globose to obovoid, 12–18 mm long, resinous. Seed
Bot. Gard. Edinburgh 45: 384. 1989; Podocarpus proper nearly spherical but slightly compressed lat-
gaussenii Woltz, Trav. Lab. Forest. Toulouse T. 1 (8, erally, 10–14 mm diam. with a verrucose surface and
2): 6. 1969. a 1 mm thick, hard seed coat.

Etymology Taxonomic notes

The species epithet refers to the (occasionally) sick- In southern Africa, this species is still classified
le-shaped leaves. under Podocarpus (see e.g. Keith Coates Palgrave’s
popular handbook Trees of Southern Africa in its
several editions, 1977–1988) despite obvious differ-
Vernacular names
ences in the lack of development of a receptacle in
Bastard yellowwood, Outeniqua yellowwood; Out- the seed cone and in the anatomy and morphology of
eniekwageelhout; inkoba (South Africa) the leaves, e.g. amphistomatic versus hypo­stomatic
leaves. Other differences are the placement of pollen
Uses
cones and seed cones below foliage leaves on scaly
dwarf shoots and the thickening and colouring of The wood of Afrocarpus falcatus, incorrectly known
the epimatium to resemble a yellow plum-like fruit. as yellowwood (= Podocarpus latifolius), is valuable
None of these characters are shared with Podo­carpus especially in the large sizes it attains in the south-
sensu stricto anywhere in the world. A form with ern Cape. It was used in the past for ship masts and
small, very narrow leaves (1.5–2.5 cm × 1.5–2 mm) is still in high demand for boat building. The sawn
from Transvaal, South Africa was described as a new timber is also used in construction for beams and
species Podocarpus gracilimus by Stapf, but there is rafters, house floors and wall panelling, carpentry
much variation in leaf sizes, which may in part be and joinery, and furniture making. In horticulture
influenced by growing conditions. The taxon origi- it is increasingly popular as an amenity tree, mainly 139
nally described as P. gaussenii from Madagascar has in countries with a mild climate like South Africa,
turned out to be a case of introduction of A. falcatus e.g. western USA, Australia and New Zealand. It can
to that island; it has not been found growing in the also be seen in several botanic gardens and arboreta
wild in recent surveys of the forest flora. outside South Africa.

Distribution
Afrocarpus gracilior (Pilg.) C. N. Page, Notes Roy.
SE & S Africa: from Malawi and Mozambique to Bot. Gard. Edinburgh 45: 383. 1989. Podocarpus
Kwazulu Natal and Eastern and Western Cape Prov- gracilior Pilg., in Engler, Pflanzenr. IV.5 [18]: 71.
inces, South Africa. 1903; Decussocarpus gracilior (Pilg.) de Laub.,
TDWG codes: 26 MLW MOZ 27 CPP-EC CPP-WC J. Arnold Arbor. 50: 359. 1969. Type: Ethiopia:
NAT OFS SWZ TVL-GA TVL-MP TVL-NP [“Gerra Abuna Tekla Zlaimanot (Haimanot), auf
Bergen 2500 m ü. M.”], G. H. W. Schimper 1160
(syntype K). Fig. 25, 26
Ecology
Afrocarpus falcatus occurs in the high, moist forests
Etymology
lining rivers in the southern Cape, where it attains its
greatest size in the Knysna forest just above sea level. The species epithet refers to the thin, slender leaves.
Several of the very large trees here may be 1000–1500
years old and the species is thus another of the large,
Vernacular names
emergent, long-lived conifers. Eastwards and north-
wards it ascends to higher elevations (alt. range 500– sigba or zigba (Amharic, Ethiopia); mponda, poda
1700 m a.s.l.) and is restricted to patches of moist (Kirangi, Tanzania)
forest in ravines and wooded slopes exposed to oce-
anic winds that bring rain. Here it attains only 25
Description
m or less. In the coastal forests it is associated with
Podocarpus latifolius, Ocotea bullata, Celtis africana, Trees to 40 m tall, d.b.h. to 2.5 m. Bark smooth
Ilex mitis, Nuxia spp. and Olea spp. and the large po- in young trees, flaking in rectangular or round-
docarp trees are usually solitary, isolated individu- ed small plates in large trees, light or dark (red-)
als or occasionally form a small group. Presumably brown, weathering grey. Branches ascending and
their regeneration is dependent on episodal canopy spreading forming a broad, domed crown. Foliage
disturbance similar to e.g. Podocarpus totara in New dense, on numerous branches; new lateral branch-
Zealand and many other big, long-lived conifers. lets ridged or more or less quadrangular; terminal
buds small, ca. 2 mm diam., or absent; bud scales
triangular with a rostrate apex. Leaves spirally ar-
Conservation
ranged, on seedlings and young plants sometimes
IUCN: LC opposite and narrowly linear-lanceolate, up to 18 cm
long and 4–8 mm wide, straight or falcate, tapering
 to a fine point. Adult leaves shorter, (1.5–)3–6(–8) TDWG codes: 24 ETH SUD 25 KEN TAN UGA
cm long, (1.5–)2–4(–5) mm wide, spirally arranged,
twisted at the narrowed base, spreading to ascend-
Ecology
ing, straight or rarely slightly falcate, linear to line-
ar-lanceolate, with a raised midrib adaxially (lower Afrocarpus gracilior occurs in montane evergreen
side), present at the proximal end or entirely absent rainforest at altitudes from 1500 m to 2600 m a.s.l.
abaxially, grey-green; apex acute. Stomata on both In a wet type of forest in Ethiopia and Kenya Olea
surfaces, arranged in numerous intermittent lines welwitschii and Afrocarpus gracilior are dominant,
not well separated by midrib. Pollen cones solitary with an understorey in which Coffea arabica is fre-
or with 2–3 on very short stalks or subsessile, axil- quent. Other dominants in similar forests elsewhere
140 lary to foliage leaves or not, subtended by a whorl are Syzygium spp., Schefflera spp., Celtis spp., Ilex
of papery bracts, initially subglobose, elongating to mitis, Ocotea kenyensis, and Nuxia congesta; bam-
cylindrical, (5–)10–18(–23) mm long, 2.5–3.5 mm boo thickets sometimes form a monotonous un-
diam.; ­microsporophylls spirally arranged, imbri- derstorey. In small forest patches interspersed with
cate before anthesis, triangular-trullate, ca. 1.5 mm subalpine grasslands (with Arundinaria alpina)
wide, with denticulate-lacerate margins and acute or Afrocarpus gracilior may occur with e.g. Albizzia sp.,
apiculate apex, bearing two subglobose pollen sacs. Cussonia holstii and Erythrina abyssinica. Many of
Seed cones solitary on small, scaly branchlets situ- these forests and forest patches have been degraded,
ated axillary to or below foliage leaves, with several but A. gracilior may still occur in the ensuing sec-
sterile and one terminal, larger fertile bract. Ma- ondary woodland vegetation, even though it is a
ture seed cones with a single seed subtended by a long-lived ‘climax’ tree. Afrocarpus gracilior also oc-
single, short bract; with seed enclosed by a fleshy, curs among or can be co-dominant with Juniperus
firm epimatium that ripens from glaucous green to procera, forming characteristic juniper-podocarp
yellow or light orange, broad ellipsoid to pyriform forests on the high plateaus and ridges. These forests
or sometimes globose, 15–20(–23) mm long. Seed occur in regions with less high rainfall than in those
proper broadly ovoid but slightly compressed lat- where angiosperms dominate.
erally, 12–18 mm long, with a smooth surface and
hard, 1–1.5 mm thick seed coat.
Conservation
IUCN: LC
Taxonomic notes
This species has been considered by some botanists
Uses
to be conspecific with Afrocarpus falcatus in South
Africa, but these species are not only geographically Afrocarpus gracilior is an important timber tree in
separated but also distinct in several, albeit minor, eastern Africa and the timber is exported as well as
morphological characters. Such distinctions can of used locally. The sawn timber is used in construc-
course seem relative; if one considers Afrocarpus and tion and particularly inside work such as floors,
Podocarpus to be one genus (as many botanists in doors and wall paneling, carpentry and joinery and
South Africa still do) then the difference between furniture making. This species has been introduced
A. falcatus and A. gracilior may appear to be less sig- from Ethiopia as a forestry plantation tree into other
nificant. The distinction between species, however, is countries, e.g. India, where trial plantations at De-
not a matter of degree, but of discontinuity of charac- hra Dun were begun in the early years of the 20th
ter states, indicating a history of genetically separated century. In northern Ethiopia it is also often planted
populations, regardless whether the two populations in church compounds to provide shade and shelter
are now geographically adjacent or disjunct. for the congregation (Prof. Ib Friis, pers. comm.). It
is not known to be introduced in horticulture and
may only be present in a few botanic gardens.
Distribution
Ethiopia, Kenya, Sudan (Equatoria), Tanzania,
Uganda.
Afrocarpus mannii (Hook. f.) C. N. Page, Notes from glaucous green to reddish brown, obliquely
Roy. Bot. Gard. Edinburgh 45: 384. 1989. Podo­ ­pyriform, 20–30(–35) mm long, resinous. Seed
carpus mannii Hook. f., J. Proc. Linn. Soc., Bot. 7: proper nearly obovoid but slightly compressed lat-
218. 1864; Nageia mannii (Hook. f.) Kuntze, Revis. erally, 16–25 mm long, with an uneven surface and a
Gen. Pl. 2: 800. 1891; Decussocarpus mannii (Hook.) 4–5 mm thick, hard seed coat.
de Laub., J. Arnold Arbor. 50: 359. 1969. Type: São
Thomé Principe: São Thomé, Pico, on the summit,
Distribution
G. Mann 1065 (holotype K).
West Central Africa (Gulf of Guinea Islands): Sao
Tomé.
Etymology
TDWG codes: 23 GGI-ST 141
This species was named after Georg Mann, who col-
lected the type specimen.
Ecology
Afrocarpus mannii is endemic on the volcano Pico
Vernacular names
de São Thomé from ca. 1450 m to the summit area
pinheiro de São Thomé, pinheiro da terra (Portu- at 2142 m a.s.l. It is nowhere a tall tree and at the
guese) summit it is reduced to dwarfed krummholz. It is
common in the high montane cloud forest where
this has remained undisturbed.
Description
Trees generally to 15 m tall but on the summit area as
Conservation
krummholz. Bark undescribed. Branches ascending
and spreading forming a broad crown. Foliage rela- Deforestation at lower to middle altitudes on the
tively sparse; new lateral branchlets ridged or more mountain is the main threat to this species, which is
or less quadrangular; terminal buds small, ca. 2 × endemic to the island.
1 mm, or absent; bud scales triangular, acuminate. IUCN: VU (D2)
Leaves spirally arranged, on seedlings and young
plants linear-lanceolate to subfalcate, up to 16 cm
Uses
long and 4–8 mm wide, straight or falcate, taper-
ing to a fine point. Adult leaves slightly shorter, (2–) The timber of Afrocarpus mannii is valuable in trees
3–8(–11) cm long, 3–7(–8) mm wide, twisted at the of good size and shape, which have become scarce. It
petiolate base, spreading, straight or slightly falcate, is used for light construction. This species has been
lanceolate to linear-lanceolate, with a conspicuously planted in rural areas in Cameroon and Ivory Coast
raised midrib adaxially (lower side) and obscurely and probably elsewhere in W Africa as a canopy tree
present abaxially, grey-green; apex acute to obtuse. or windbreak for coffee plantations and as an amen-
Stomata on both surfaces, arranged in numerous ity tree in villages.
intermittent lines not well separated by the midrib.
Pollen cones solitary or with 2, sessile, axillary to fo-
liage leaves or not, subtended by a few scaly bracts, Afrocarpus usambarensis (Pilg.) C. N. Page,
initially subglobose, elongating to cylindrical, 10–20 Notes Roy. Bot. Gard. Edinburgh 45: 384. 1989.
mm long, 2–3 mm diam.; microsporophylls spirally Podocarpus usambarensis Pilg., in Engler, Pflanzenr.
arranged, imbricate before anthesis, broadly trian- IV.5 [18]: 70. 1903. Type: Tanzania: Tanga Prov.,
gular, ca. 1 × 1 mm, with lacerate margins and acute Usambara Mts., Mtai Hill, near Mpare village,
apex, bearing two pollen sacs. Seed cones solitary C. Holst 2467 (syntype K).
on small, scaly branchlets situated axillary to foliage
leaves, with several sterile bracts and one terminal,
Etymology
larger fertile bract. Mature seed cones with a single
seed subtended by a single, short bract; with seed The species epithet refers to the Usambara Moun-
enclosed by a fleshy, firm epimatium that ripens tains from where it was first described.
 Vernacular names Ecology
mse, muze (Usambara Mts., Tanzania) Afrocarpus usambarensis occurs in montane ever-
green rainforest and dry evergreen forest, mixed
with co-dominant angiosperms. Elevation ranges
Description
from ca. 1500 m to ca. 3000 m. a.s.l. Trees are of-
Trees to 30 m tall, d.b.h. to 2 m. Bark smooth in ten solitary but not emergent, only reaching into the
young trees, flaking in rectangular or rounded general canopy of the forest. In rainforest it occurs
small plates in large trees, dark brown, weathering often with Podocarpus milanjianus; the co-dominant
grey. Branches ascending and spreading forming a angiosperm tree in this wetter forest type is often
142 broad, domed crown. Foliage dense, on numerous Ocotea usambarensis, but many other species may
branches; new lateral branchlets ridged or more or occur with it. In dryer evergreen forest Olea and
less quadrangular; terminal buds very small, 0.6–1 ­Ficus are common associates of A. usambarensis,
mm diam., or absent; bud scales rounded with or other taxa are e.g. Calodendrum capense, ­Syzygium
without an apiculate apex. Leaves on seedlings cordatum and Bridelia micrantha. These drier forests
and young plants mostly opposite, narrowly line- are often degraded or converted to coarse grassland
ar-lanceolate, up to 13 cm long and 4–7 mm wide, in which A. usambarensis can survive as isolated
straight or falcate, tapering to a fine point. Adult trees, at least for a time.
leaves shorter, (1.5–)3–5(–6) cm long, (1.5–)2–4(–5)
mm wide, spirally arranged, twisted at the nar-
Conservation
rowed base, spreading to ascending, straight or
rarely slightly falcate, linear-elliptic or with paral- This species is considered Vulnerable based on di-
lel sides, gradually tapered above 2/3 or 3/4 of their rect observation of intense local exploitation (aer-
length, with a raised midrib usually present on both ial photography of many saw pits both within and
sides, grey-green; apex acute. Stomata on both sur- without reserves). This species is under severe threat
faces, arranged in numerous intermittent lines not from illegal logging in the Chome Forest Reserve in
well separated by the midrib. Pollen cones solitary Tanzania (evidence from aerial photography); the
or with 2–3 on very short stalks or subsessile, axil- same type of saw pit exploitation is known from
lary to foliage leaves or not, subtended by a whorl other locations. General deforestation and fires are
of papery bracts, initially subglobose, elongating to also reducing the rainforest, which is usually lim-
cylindrical, (5–)10–20(–26) mm long, 2.5–3.5 mm ited in extend even naturally. This species is the
diam.; microsporophylls spirally arranged, imbri- most valuable and specifically targeted tree for (il-
cate before anthesis, triangular-trullate, ca. 0.8 mm legal) logging in this type of forest. It is present in
wide, with denticulate-lacerate margins and acute or the following reserves in Tanzania: Chome, Hanang,
apiculate apex, bearing two subglobose pollen sacs. Mafwomero, Mkusu, Nou, Shagayu and Wotta. The
Seed cones solitary on small, scaly or leafy branch- evidence is that this does not prevent illegal logging
lets situated axillary to or below foliage leaves, with on a large scale using sawpits to remove and process
several sterile and one terminal, larger fertile bract. individual trees.
Mature seed cones with a single seed subtended by IUCN: VU (A2, c, d)
a single, short bract; seed enclosed by a fleshy, firm
epimatium that ripens from green or glaucous green
Uses
to yellow, globose or sometimes broad ellipsoid,
(17–)23–30(–35) mm long. Seed proper globose or This species, yielding ‘yellowwood’ or podocarp
broadly ellipsoid but slightly compressed laterally, wood is highly valued for its timber and exploited
(15–)20–25(–30) mm long, with a rugose-pusticulate mainly for sawn timber used in construction of
surface and a (3–)4–6(–8) mm thick, hard seed coat. houses. The wood is yellowish in colour, straight-
grained, and clean of knots and can be used for gen-
eral carpentry and furniture as well. This species is
Distribution
not known to be in cultivation.
Tanzania, Kenya (Kyulu Hills, Taita Taveta District).
TDWG codes: 25 KEN TAN
Agathis Salisb., Trans. Linn. Soc. London 8: 311. 1807 (nom. cons.). Dammara
(Rumph.) Lam., Enum. Pl. Hort. Berol. 2: 411. 1822. Type: Agathis dammara
(Lamb.) Rich. & A. Rich. [Dammara loranthifolia Link (Pinus dammara Lamb.)]
(Araucariaceae).

Salisburyodendron A. V. Bobrov & Melikyan, Koma­ and rudimentary to sometimes virtually absent.
rovia 4: 62. 2006. Type: Salisburyodendron australis Seedlings with 2 cotyledons.
(D. Don) A. V. Bobrov & Melikyan [Agathis australis
(D. Don) Lindl.]. 17 species. 143

Greek Agathis = a clew or ball of thread; it refers to

Distribution
the seed cone.
Malesia: Malay Peninsula, Sumatera, Borneo, Su-
lawesi, Philippines, Maluku [Moluccas], New Guin-
Description
ea, New Britain; Australia: coastal Queensland;
Evergreen monoecious trees, often of great size, mo- SW Pacific: New Caledonia, Vanuatu, Fiji, Solomon
nopodial with straight boles. Resin canals in bark, Islands (Santa Cruz Group), New Zealand (North
leaves and seed cones. Branching in sub-verticillate Island).
pseudo-whorls (Massart’s or Rauh’s model), trun-
cated in very large trees. Apical buds globose with
Taxonomic notes
­imbricate scales. Leaves subopposite to opposite,
short petiolate, broad and multi-veined, coriaceous, Few attempts have been made to investigate the pos-
more or less hypostomatic, extremely variable in sible relationships among the various species in this
shape and size within a single tree; those on young genus. Most studies of this kind have concerned
trees usually larger than on mature trees. Pollen themselves with relationships at the level of genus.
cones appearing after seed cones, axillary, solitary, Phylogenetic relationships of taxa within the Arau-
sessile to pedunculate, subtended by more or less cariaceae were investigated by Gilmore & Hill (1997),
decussate bracts, the lower pair of which may be Setoguchi et al. (1998) and Kershaw & Wagstaff
leaf-like or not, elongating to a cylindrical or catkin- (2001) (the first two papers used rbcL DNA mark-
like shape after anthesis. Microsporophylls helically ers) and none of these studies sampled all species.
attached to a rachis in imbricate or tesselate arrange- We therefore still lack a hypothesis of the phylogeny
ment, consisting of a short stalk and a more or less of the species of Agathis upon which a classification
peltate head of varied shape according to species; of the genus could be based and, indeed, no infra­
bearing from 2–12 elongated pollen sacs directed generic classification has been proposed. In order to
inward towards cone rachis. Seed cones axillary or conveniently and more reliably key out the species,
sometimes terminal, solitary on stout stalks, globose they have here been divided into two geographically
to obovoid, usually smooth but sometimes rough determined groups; each of these covers roughly
with bossed scale margins, green or slightly glau- half of the entire range of the genus. No taxonomy is
cous, disintegrating when mature and drying. Cone implied with this division and it only serves the keys.
scales composed of a fused bract and seed scale, the The South-West Pacific here includes New Zealand
visible and largest part made up of the bractaceous and Malesia is the region, as defined in Flora Male-
element, helically attached to a stout rachis short- siana, from Peninsular Malaysia to New Guinea and
er than the cone, imbricate, thin except the outer, the Solomon Islands. Few species are known in cul-
mostly exposed margin. Seeds 1 per cone scale, in- tivation outside their ‘own’ region as here defined
verted, more or less ovoid but strongly flattened, (the exceptions almost all involve A. australis from
with two thin, membranous wings, one obliquely New Zealand) and a key to cultivated species is for
placed relative to the seed axis, the other opposite this reason not considered necessary.
 6b. Pollen cones with 6–16 decussate, imbricate or
Key to the species of Agathis in Australia and
free, linear to triangular bract scales. Juvenile
the SW Pacific
leaves 11–20 cm long; adult leaves (2.5–)4–9 cm
Pollen cones provide most of the diagnostic char- long 7
acters of species in this genus; under mature trees 7a. Pollen cones short cylindrical to nearly oval,
(all are monoecious) these are often found in the leaf 2–2.5 cm long, 8–10 mm wide; intact cones
litter under their canopy. Only fully expanded cones with 2 leaf-like bracts subtending 6–8 free bract
should be taken into account. Leaves are highly scales A. lanceolata
variable but differ between ‘juvenile’ (phase 1) leaves 7b. Pollen cones cylindrical, (2–)2.5–5(–7) cm long,
and ‘adult’ (phase 2) leaves as treated in the species 6–15 mm wide; lacking 2 leaf-like bracts, with
144 descriptions; only maximum length for the first cat- 8–16 imbricate bract scales 8
egory appears to be informative in this context. A 8a. Bark on trunk with numerous small, granular
single twig with foliage can never be determined lenticels. Bract scales at base of pollen cone 1–2
with the macroscopic leaf characters (measure- mm wide; microsporophyll heads umbonate.
ments and shapes) alone. Seed cones up to 7 cm diam A. montana
8b. Bark on trunk without or with few lenticels.
1a. Pollen cones short cylindrical, 0.9–1.6 cm long, Bract scales at base of pollen cone 4–7 mm
4–8 mm wide 2 wide; microsporophyll heads nearly flat or con-
1b. Pollen cones cylindrical or oval-fusiform, vex. Seed cones 9–13 cm diam 9
2–6(–7) cm long, (6–)7–18 mm wide 3 9a. Pollen cones with 8–16 decussate bract scales
2a. Microsporophylls of pollen cones imbricate, at base, 6–9 mm wide; microsporophyll heads
more or less convex. Seed cones 3.5–5 cm diam.; nearly flat with angular upper margin
cone scales with a bossed upper margin A. moorei
A. atropurpurea 9b. Pollen cones with up to 8 decussate bracts
2b. Microsporophylls of pollen cones tessellate, scales at base, 8–15 mm wide; microsporophyll
prismatic. Seed cones 6.5–10 cm diam.; cone heads convex, often notched at apex
scales with a smooth, slightly rounded upper A. macrophylla
margin A. microstachya
3a. Juvenile leaves to 6 cm long. Pollen cones on
Key to the species of Agathis in Malesia
(4–)5–20 mm long, stout peduncles; basal bract
scales short, rounded or triangular 5 1a. Juvenile (phase 1) leaves to 8 cm long, adult
3b. Juvenile leaves to 20 cm long; pollen cones ses- (phase 2) leaves to 5 cm (occasionally to 6–7
sile or on short peduncles to 7 mm long; basal cm) long, the shortest leaves often obovate or
bract scales 1–7 mm wide, linear to triangular oval-orbiculate (length less than 2 × width) 2
4 1b. Juvenile (phase 1) leaves to 14 cm long, adult
4a. Bark exfoliating with small flakes (smaller than (phase 2) leaves to 12(–14) cm long, the shortest
10 cm). Pollen cones 8–13 mm wide, with 8–10 leaves variable in shape but not obovate or oval-
decussate, imbricate basal bract scales orbiculate 4
A. robusta 2a. Pollen cones when fully expanded 1.2–1.5 cm
4b. Bark exfoliating with large flakes (to 15 cm). long, 5–6 mm wide, lacking 2 leaf-like bracts
Pollen cones 15–18 mm wide, with 4–6 decus- subtending the cluster of smaller bract scales
sate, free basal bract scales A. silbae A. orbicula
5a. Bark on trunk smooth, with small or large 2b. Pollen cones when fully expanded 2.5–4 cm
flakes. Trees becoming very large 6 long, 8–10 mm wide; intact cones usually with
5b. Bark on trunk fissured, rough and scaly. Trees 2 leaf-like bracts up to 3 cm long, subtending a
often stunted, up to 10 m tall (rarely taller to cluster of smaller bract scales 3
25 m in forest) A. ovata 3a. Two subtending bracts of pollen cones to 30 ×
6a. Pollen cones with 6 decussate, imbricate, short 10 mm. Seed cone scales with a boss on the up-
rounded bracts scales. Juvenile leaves to 6 cm per margin. Endemic of Peninsular Malaysia
long; adult leaves 2.3–4(–7) cm long A. flavescens
A. australis
3b. Two subtending bracts of pollen cones to 20 ×
Description
4 mm. Seed cone scales lacking a boss on the
upper margin. Endemic of Borneo Trees to 50 m tall and 2.5 m or more d.b.h. with
A. kinabaluensis clear cylindrical bole to 30 m and a rounded or
4a. Pollen cones when fully expanded 8–9 cm long, open crown with spreading to ascending branches.
25–35 mm wide; short basal bracts in 2 decus- Bark smooth or sometimes scaly, exfoliating in ir-
sate pairs; microsporophyll heads 5–6 × 4–8 regular thin scales of up to 15 cm across; outer bark
mm (length × width) A. borneensis dark brown or red-brown, with purple patches un-
4b. Pollen cones when fully expanded (2.2–)3–6 der newly fallen scales; inner bark reddish, exuding
(–7) cm long, 8–13 mm wide; short basal bracts white resin. Leaves subopposite, thin or thick, multi-
in 2–5 decussate pairs; microsporophyll heads nerved, coriaceous, glabrous, light green, sometimes 145
0.7–2 × 1–2.5 mm (length × width) 5 glaucous on the underside especially leaves on new
5a. Microsporophylls in tessellate arrangement, shoots. Leaves on saplings and young trees and/or in
their heads prismatic; short basal bracts of pol- shade linear-lanceolate to elliptic, 5–8 cm long, 1.5–3
len cones in 4–5 decussate pairs 6 cm wide; leaves in crowns of mature trees lanceolate
5b. Microsporophylls in imbricate arrangement, to elliptic, 3–7 cm long, (0.5–)1–2 cm wide, short pet-
their heads more or less convex; short basal iolate and with an obtuse apex. Pollen cones axillary,
bracts of pollen cones in 2–4 decussate pairs 7 solitary on a 2–3 mm long, stout peduncle, short cy-
6a. Pollen cones when fully expanded barrel- lindrical, when full grown 0.9–1.6 cm long, 4–7.5 mm
shaped, 2.2–3 cm long, 10–12 mm wide. Bark diam., with 8–10 decussate (in 4–5 pairs), imbricate,
on large trunks with large (to 10 cm) flakes 1–2.5 mm wide bract scales at their base. Microsporo-
A. labillardierei phylls in imbricate arrangement; head with rounded
6b. Pollen cones when fully expanded cylindrical, or minutely mucronate upper margin, 0.6–0.8 mm
4–6(–7) cm long, 8–13 mm wide. Bark on large wide, 0.5–0.7 mm high in mature cones, bearing
trunks with small (to 5 cm) flakes A. robusta 2–5 pollen sacs. Seed cones solitary on thick pedun-
7a. Short basal bracts of pollen cones in 2–4 dec- cles, globose, 3.5–5.5 cm long, 3.5–5 cm wide, more
ussate pairs, free spreading; peduncle absent or or less rough, green and resinous, ripening brown.
to 5 mm long. Seed cone scales with smooth, Cone scales with thick, slightly bossed, incurved up-
slightly rounded margins A. dammara per margins, 1.6–2.3 cm long, 2.3–3 cm wide, broad
7b. Short basal bracts of pollen cones in 2–3 decus- triangular to reniform with rounded corners and
sate pairs, imbricate; peduncle 2–10 mm long. more or less flanged on either side. Seeds 7–9 × 4–5
Seed cone scales with bossed upper margin mm, ovoid, flattened, with two unequal wings; larg-
A. lenticula est wing 12–15 × 8 mm; smallest wing a small acute
triangular point 2 mm long opposite largest wing.

Agathis atropurpurea B. Hyland, Brunonia

Distribution
1 (1): 109. 1978. Type: Australia: Queensland,
Cook District, Bellenden Ker, [“N. P. R. 226”], Australia: NE Queensland (Cook District).
B. P. M. Hyland 5776 (holotype BRI). TDWG codes: 50 QLD-QU

Etymology Ecology
The species epithet describes the colour of the bark Agathis atropurpurea is a rare species occurring in
as dark or blackish purple (Latin atratus = dark, lower montane rainforest at altitudes between 900
blackish; purpureus = purple). m and 1500 m a.s.l. on granite mountains and out-
crops which rise above the Atherton Tableland.
Vernacular names
Conservation
Blue kauri pine, Black kauri pine
Wherever logging is/was allowed exploitation has
been heavy and subpopulations have declined as
 a result. The extent of the decline is not accurately opposite, thick, multinerved, coriaceous, glabrous,
recorded; if this were better quantified this species frequently glaucous to farinose when young, espe-
could meet criterion A1c,d under Vulnerable (IUCN cially on the abaxial (lower) surface. Leaves on sap-
Red List Categories and Criteria version 3.1, 2001). lings and young or shaded trees variable in size and
A large percentage of the remaining forest is now shape, usually lanceolate, 3.5–6 cm long, 6–10 mm
protected. Part of the population of A. atropurpurea wide, with acute apex, but occasionally much larger
is protected in Wurunuru National Park. (9 cm long, 3.5 cm wide) and with rounded apices.
IUCN: NT Leaves in crowns of mature trees much smaller, 2.3–
4 (rarely 7) cm long, 9–15 mm wide, broadly ovate
in shape, often with a truncate apex. Pollen cones
Uses
146 axillary, solitary on a stout , 5–15 mm long pedun-
Blue kauri pine used to be a valuable timber tree cle, cylindrical, when fully mature 3–5 cm long, 7–10
in the days of logging the old growth forests of mm diam., ripening from yellowish green to brown;
Queensland. The large dimensions of these trees, basal bract cluster loose, usually wider than pollen
combined with the excellent properties for carpen- cone, typically consisting of three opposite pairs of
try and veneer work such as light weight, even and imbricate, short, rounded bracts and frequently with
fine grain and texture with hardly visible growth up to two long-bracts 15–20 × 3–5 mm immediately
rings and light colour, made them desirable. Today, subtending the basal bract cluster. Microsporophylls
few trees are logged from natural stands, as most are imbricate; pedicels 2.2–2.6 mm long, slender; heads
now protected. This species is uncommonly planted 1.8–2.5 mm wide, 2.1–2.7 mm high in cone, in adax-
as an ornamental tree and no commercial timber ial view thick with narrow marginal flange; margins
plantations of it exist. entire or rarely minutely erose; apex often notched.
Seed cones solitary on thick peduncles, sometimes
2–4 cones together on a branch, globose, 5–7.5 cm
Agathis australis (D. Don) Lindl., in Loudon, diam., glaucous white-green, green or blue-green,
Encycl. Pl.: 802. 1829. Dammara australis D. Don, (very) resinous, ripening brown. Cone scales im-
in Lambert, Descr. Pinus 2: 14, t. 6. 1824; Salis­ bricate, spreading distally at pollination time, later
buryodendron australis (D. Don.) A. V. Bobrov closing and becoming rhombic-rostellate, giving
& Melikyan, Komarovia 4: 63. 2006. Type not mature seed cones a rough, scaly surface, ca 2 cm
designated. Fig. 27, 28, 29 long, 3 cm wide. Seeds 8–10 × 5 mm, ovoid-cuneate,
with two unequal wings; largest wing ca. 12 × 8 mm,
more or less square; smallest wing rudimentiary or
Etymology
virtually absent.
The species epithet means ‘from the south’ and prob-
ably refers to the fact that this is the southernmost
Distribution
species in the genus.
New Zealand: North Island (Northland).
TDWG codes: 51 NZN
Vernacular names
Kauri pine; kauri (Maori)
Ecology
Agathis australis is the dominant tree in mixed con-
Description
ifer-angiosperm subtropical lowland primary forest
Trees to 55 m tall, to 5 m d.b.h. or more in veteran that once covered the northern peninsulas of North
trees, self-pruning and with a long, clear, almost cy- Island from near sea level to 375 (–600?) m a.s.l., but
lindrical bole. Bark smooth, peeling frequently in of which only small remnants remain today. It is not
irregular small patches, often creating a mix of col- clear why it did not occur naturally further south
ours on a single tree from salmon-pink through to on the island, as planted trees grow well there now.
grey. Branches in young trees semi-whorled, spread- Perhaps the dynamics of the Kauri forest in its natu-
ing, in large trees ascending from the top of the ral state played a part in this limitation. The species
bole, forming a broad, domed crown. Leaves sub- is dependent for successful regeneration on episodal
disturbance of the forest by fire or storm removing tralis would fulfil the criteria for a listing as (at least)
sizable swathes of forest cover. Even-aged stands of Endangered (EN), even though the causes of decline
large trees in groves and a few scattered giant trees have now ceased, because the reduction amounts to
are a common pattern in the Waipoua Forest, the more than 70% over the last three generations (of
largest of the remaining primeaval Kauri forest mature trees). Conservationists in New Zealand ob-
remnants. The giants are more than 1000 years old ject to such a rating because to them it would appear
and have survived one or more disturbances, act- to be a denial of their successful efforts to curb the
ing as seed trees. In such a mature stand of Kauri destruction and save this iconic tree from extinc-
there is little regeneration. Forest succession with- tion. On the other hand one could observe that most
out disturbance eventually leads to dominance of of the historical reduction will be permanent under
angiosperms. Early phases in disturbed areas are human occupation and land use and that while the 147
dominated by Leptospermum scoparium and Kunzea reduction has ceased (and is even being reversed)
ericoides (Myrtaceae) into which A. australis invades no one can guarantee what the priorities of future
abundantly. The conifer Phyllocladus trichomanoides generations are going to be.
arrives next. Via stages with increased tree species IUCN: LC
diversity and the establishment of more shade toler-
ant trees, above which the ‘rickers’, i.e. Kauri trees
Uses
with conical habit, rise, the forest restores itself.
A drastic thinning of Kauri trees then results in The kauri of New Zealand was once the most impor-
fewer trees with ever wider spreading crowns, ris- tant timber tree of these southern islands, but in the
ing above the canopy as emergents. Eventually there short space of roughly 50 years this natural resource
is another disturbance, but these cycles may have a had nearly been exhausted through one of the most
duration of 500 years or longer. Common conifers wanton campaigns of exploitation and short-term
in mature Kauri stands are Dacrydium cupressinum, thinking in the history of European colonisation of
Prumnopitys ferruginea, P. taxifolia, Podocarpus foreign lands. For several decades, the protection of
totara and Dacrycarpus dacrydioides; common co- remaining forests and regeneration projects have
dominant angiosperms are e.g. Beilschmiedia spp., now ensured its continuing existence, but the great
Weinmannia racemosa and Metrosideros umbellata. expanse of kauri forest has nevertheless been greatly
reduced. The timber of old growth stands is/was of
large dimensions and the properties of the wood are
Conservation
excellent for joiner’s work, boat building and car-
Since the arrival of Europeans in New Zealand, the pentry. ‘Black kauri’, dark brown in colour and very
estimated 1,215,000 ha once covered by ­primeaval hard and durable, came from logs buried in peat de-
kauri forests have been reduced to ca. 7,500 ha. posits. Wood from burls is beautifully figured and
These remnants are now strictly protected; the larg- was used in furniture and for panelling. The vast
est reserve is the Waipoua Forest covering 9,105 ha abundance of timber in the heyday of exploitation
(not all of it is dominated by A. australis) created meant that it was also put to less refined use, such as
in 1952. However, natural regeneration, where land mine-props and railway sleepers. Another valuable
use as mixed (semi-)natural forest has not changed, product is resin (copal) of which this tree produces
is abundant in many places because the species of great quantities. During and after the timber exploi-
course does not distinguish between natural and tation, a veritable ‘copal rush’ took place, with thou-
man-made disturbance events. This regeneration sands of resin diggers coming from all over to dig up
is possible, in ecologically suitable locations, with- the semi-fossil resin in the cut-over forests. Much of
in some 80,000 ha of existing forest reserves. The this was used in the manufacture of linoleum, paint
Department of Conservation in New Zealand has and varnish. This species has been taken into cul-
plans to establish a Kauri National Park in order to tivation both in (modest scale) forestry plantations
give full protection to these forests. If IUCN Red and as an ornamental tree in several countries with
List criteria were strictly applied to this history of a mild climate.
reduction of population size (A criterion), A. aus-
 Agathis borneensis Warb., Monsunia 1: 184. 1900. brown. Cone scales with slightly rounded, incurved
Type: Malaysia: Sarawak, [locality not stated], or slightly projecting upper margins, ca. 3.5 cm long,
O. Beccari 596 (syntype K). Pl. 5, Fig. 30 3.5–4.5 cm wide in larger cones, roughly triangular
with rounded corners and more or less flanged on
Agathis endertii Meijer Drees, Bull. Jard. Bot. Buiten- either side. Seeds 12–15 × 7–8 mm, ovoid-oblong,
zorg, ser. 3, 16: 470. 1940. with two unequal wings; largest wing ca. 20 × 13
mm; smallest wing a small blunt triangle 3–5 mm
wide opposite largest wing.
Etymology
The species epithet refers to Borneo, where it is
Taxonomic notes
148 ­native.
Agathis borneensis has been confused with A. dam-
mara in the past, which is perhaps understandable
Vernacular names
since the diagnostic characters that distinguish the
Numerous local common names are applied to this two species are often not present even in quite large
species. Some of these are general names for Agathis trees. These are mainly found in the pollen cones,
(like kauri, the Maori word adopted in English) and which can often been found underneath trees on
I cite only a few: bindang (Brunei, Sarawak); bem- the forest floor but can decay quickly in the tropical
bueng (Kalimantan) damar minyak (Malay Penin- climate. The cones of A. borneensis are more robust
sula); damar pilau (Dayak, Kalimantan); tambunan and more or less globose to oblong until elongation
(Sabah); hedje (Sumatera). at anthesis makes them cylindrical; they are more
cylindrical from the start in A. dammara. The mi-
crosporophylls of A. borneensis are much larger than
Description
those of A. dammara and have a distinct, lighter
Trees to 50(–55) m tall and 3.5 m or more d.b.h. with and thin upper margin. When using these charac-
clear cylindrical bole up to 20–30 m and a broad ters the two species are found to be geographically
crown radiating above. Bark variable; smooth or separated. Agathis endertii Meijer Drees is listed
dippled and lenticellate or very rough and scaly, col- as an accepted species in the World Checklist and
our grey, dark brown or blackish outside, reddish Bibliography of Conifers (Farjon, 1998, [2001]). De
brown or yellowish brown under outer, exfoliating Laubenfels (1988) placed it in a section separate
layers. Leaves subopposite, thick, multinerved, co- from A. borneensis based on the shape of scales in
riaceous, glabrous, light green. Leaves on saplings seed cones (projected apex), but it is not really dis-
and young trees and/or in shade lanceolate to (nar- tinct in any diagnostic character and, in agreement
rowly) elliptic, up to 14 cm long, 1.5–4 cm wide, of- with Whitmore (1980) is here treated as a synonym
ten acute; leaves in crowns of mature trees distinctly of A. borneensis.
petiolate; shape variable, ovate to ovate-elliptic or
sometimes (narrowly) lanceolate, 2.5–10 cm long,
Distribution
(1–)2–5 cm wide, mostly with an obtuse apex. Pol-
len cones axillary, solitary on a 1–5(–8) mm long, Borneo, Malay Peninsula, Sumatera.
stout peduncle, when immature from globose be- TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
coming nearly cylindrical, 2–4 cm long, 10–25 mm MLY-PM SUM
diam., elongating past anthesis to 8–9 cm long and
up to 35 mm wide, with 4 decussate (in 2 pairs) 5–8
Ecology
mm wide, free spreading bract scales at their base.
Microsporophylls in imbricate arrangement; head Agathis borneensis occurs in lowland to upland trop-
slightly convex towards the upper, rounded, paler ical rainforest as scattered emergent trees and in low
coloured, minutely erose-denticulate margin, 4–8 lying kerangas forest on sandy or sometimes peaty
mm wide and 5–6 mm high in mature cones, bear- soils, where it can form extensive pure stands, or oc-
ing 4–10 oblong pollen sacs. Seed cones solitary on curs mixed with the following conifers: Dacrydium
thick peduncles, broadly ellipsoid to globose, to pectinatum, Falcatifolium falciforme, Nageia walli-
10–13 cm diam., smooth, green, resinous, ripening chiana, Podocarpus spp., and Sundacarpus amarus.
 149

3
4

Pl ate 5. Agathis borneensis. 1. Habit of tree. 2. Foliage branch. 3. Leaf. 4. Pollen cone. 5. Microsporo-
phylls. 6. Immature pollen cone. 7. Seed cone.
 The most common angiosperm tree on peaty soils a trend to shift from round wood to sawn timber,
growing with A. borneensis is probably Gonystylus which fetches much higher prizes. This species (and
bancanus (Thymaelaeaceae). In lowland to lower A. dammara) are planted on a fairly large scale in for-
montane rainforest it can be associated with Dipte- estry plantations in Jawa, but only locally on a small
rocarpaceae and/or Fagaceae; however, Agathis often scale within its native range. This will have to change
retreats to ridges with thin, rocky soils or to water- dramatically if the resource is to be made any­where
logged areas where these dominant angiosperms are near sustainable for the future. Agathis borneensis is
less vigorous. It is also reported from “heath for- present in some tropical botanic gardens.
est” which occurs on higher mountain ridges and
summits and is usually dominated by species in the
150 Myrtaceae. The altitudinal range of A. borneensis is Agathis dammara (Lamb.) Rich. & A. Rich., in
substantial, from near sea level to ca. 2400 m a.s.l., A. Richard (ed.) Comm. Bot. Conif. Cycad.: 83.
but with greater abuncance below ca. 1200 m a.s.l. 1826. Pinus dammara Lamb., Descr. Pinus 1: 61,
t. 38. 1803. Type: Illustration in Rumphius, Herb.
Amboinense 2: 174, t. 57. 1741 (lectotype).
Conservation
This species has been very heavily over-exploited in Agathis celebica (Koord.) Warb., Monsunia 1: 185.
many areas and as a result its total area of occupancy 1900; Dammara celebica Koord., Meded. Lands
(AOO) is estimated to have at least been reduced by Plantentuin 19: 263. 1898.
half and this is still ongoing. Stands covering an es- Agathis philippinensis Warb., Monsunia 1: 185. 1900.
timated total of 30,000 ha discovered in Kalimantan
in the 1930s had effectively been logged out by the
Etymology
mid 1960s. Most stands outside the few well pro-
tected nature reserves (mostly situated in the Malay ’Dammar’ is the local (Moluccan) name for the res-
­Peninsula and in Sabah) have been seriously deplet- in, both subfossil (‘copal’) and recent, of this tree.
ed and it is doubted that regeneration will be suf-
ficient to restore the losses. Habitat degradation has
Vernacular names
caused further reductions in recruitment of young
trees to replace felled ones. Amboina pitch tree; dammar raja (Indonesia); dam-
IUCN: EN (A4cd) mar malolo, dammar lulu (Sulawesi); almaciga,
saleng (Philippines); kaláne, kèssi, oeneëla (Maluku
[Moluccas]) and many other local names (see e.g. in
Uses
Flora Malesiana, ser. 1, 10 (3): 438, 1988).
This species is one of the most valuable and sought
after timber trees in Southeast Asia. It produces
Description
lightweight, almost white to pale yellowish wood
with no visible growth rings and a very fine and even Trees to 55(–65) m tall and 3.5–4 m or more d.b.h.
texture and without resin. It is not durable, so it will with clear cylindrical bole up to 20–25 m and a
mostly find indoor uses, but these are many, from broad crown radiating above. Bark variable; smooth
light construction and carpentry, joinery, masts for or dippled and lenticellate or rough and scaly, col-
boats, and panelling to veneer and tool or furniture our grey, dark brown or blackish outside, reddish
making. Drawing boards are made of its wood as it brown or yellowish brown under outer, exfoliating
is extremely easy to plain to a smooth surface. Its layers. Leaves subopposite, thick, multinerved, co-
odourless quality was noted in the manufacture of riaceous, glabrous, light green. Leaves on saplings
food containers, until plastics took over from it. and young trees and/or in shade lanceolate to (nar-
The inner bark exudes a translucent to white resin rowly) elliptic, up to 14 cm long, 1.5–4 cm wide,
known as ‘copal’ and is still used for varnishes in ­often acute, sometimes acuminate; leaves in crowns
photographic colour prints and as a component for of mature trees distinctly petiolate; shape variable,
the paint used to make lines etc. on tarmack road ovate to ovate-elliptic or sometimes (narrowly) lan-
surfaces. There is still a large export trade in its tim- ceolate, 2.5–8(–9) cm long, (1–)2–4 cm wide, with an
ber, but with a deminishing quantity per annum and obtuse or acute apex. Pollen cones axillary, ­solitary
on a 0–5 mm long, stout peduncle, when imma- species. This is not so with the trio A. celebica,
ture a small cylinder, 1–2 cm long, 6–8 mm diam., A. philippinensis and A. dammara; the distinctions
elongating past anthesis to 3–4 cm long and up to mentioned in Flora Malesiana (De Laubenfels, 1988)
12 mm wide, with 4–8 decussate (in 2–4 pairs) 2–4 e.g. acuminate juvenile leaves (meaning: leaves on
mm wide, free spreading bract scales at their base. juvenile trees) are found occasionally in specimens
Microsporophylls in imbricate arrangement; head from all (major) Malesian islands and are clearly
slightly convex towards the upper, rounded or re- just one of the possible shapes of these highly vari-
tuse, minutely erose-denticulate margin, 2–2.5 mm able leaves. The diagnostic pollen cones are a much
wide and 1–1.5 mm high in mature cones, bearing better organ than the leaves on young trees to look
3–6 oblong pollen sacs. Seed cones solitary on thick for consistent characters, and they unite A. celebica,
peduncles, broadly ellipsoid to globose, to 10–13 cm A. dammara and A. philippinensis, a species which 151
diam., smooth, green, resinous, ripening brown. therefore bears the earliest name. Other characters,
Cone scales with slightly rounded, incurved upper some involving leaves of mature trees, do separate
margins, ca. 3.5 cm long, 3.5–4.5 cm wide in larger some of the montane populations in Borneo, and are
cones, roughly triangular with rounded corners and here not sunk into A. dammara.
more or less flanged on either side. Seeds 12–15 × 7–8
mm, ovoid-oblong, with two unequal wings; largest
Distribution
wing ca. 20 × 13 mm; smallest wing a small blunt
triangle 3–5 mm wide opposite largest wing. Malesia: Maluku [Moluccas], Philippines, Sulawesi.
TDWG codes: 42 MOL PHI SUL
Taxonomic notes
Ecology
The nomenclature of this species is complicated,
reflecting in part conflicting views on its taxonomy Agathis dammara occurs in lowland to upland tropi-
for nearly 200 years. In more recent time, Whitmore cal rainforest as scattered emergent trees. In lowland
(1980) included not only all trees belonging to the to lower montane rainforest it can be associated with
genus occurring in Sulawesi, the Moluccas and the Dipterocarpaceae and/or Fagaceae; however, Agathis
Philippines, but also several montane populations in often retreats to ridges with thin, rocky soils or to
Borneo and the Malay Peninsula in A. dammara. De water-logged areas where these dominant angio­
Laubenfels (1988) considered A. philippinensis a dis- sperms are less vigorous. The species occurs on a
tinct species, occurring in the Philippines but also wide variety of substrates, from white sand to peaty
in Sulawesi and on the Moluccas. He furthermore soils, volcanic soils, metamorphic rock such as ser-
recognized A. celebica in Sulawesi and the Moluc- pentine or schist, or limestone. The altitudinal range
cas, with a few outliers in the Philippines. In addi- of A. dammara is from near sea level to ca. 2200 m
tion, some of the montane populations cited above a.s.l., but with greater abuncance below ca. 1200 m
(Whitmore, 1980) were by De Laubenfels separated a.s.l.
as distinct species. De Laubenfels (1988) complicat-
ed the nomenclatural knot by asserting that, under
Conservation
the rules of ICBN and in case a proposal to reject the
name Pinus dammara Lamb. in favour of A. borne­ This species has been over-exploited in many areas
ensis Warb. was not accepted at the next Interna- and as a result its total area of occupancy (AOO) is es-
tional Botanical Congress, A. borneensis would have timated to have at least been reduced by 30% or more
to be called A. dammara. His taxonomic views were and this is still ongoing. The tapping of resin when
followed in the World Checklist & Bibliography of exploited too intensively has killed large numbers
Conifers (Farjon, 1998, [2001]). This proposal was of trees in the forests, especially in the Philippines.
indeed rejected, but only if one sinks A. borneensis Habitat degradation has caused further reductions
(the western species) taxonomically into A. dam- in recruitment of young trees to replace felled ones.
mara (the eastern species) would the resulting taxon There is now a total ban on cutting Agathis trees in
have to bear that name. The type of A. dammara is the remaining forests in the Philippines, but there is
from Amboina, not from Borneo, and the pollen still illegal logging going on in some ­areas.
cones of the two are very different. These are ­separate IUCN: VU (A4cd)
 pled and lenticellate, colour grey, reddish brown
Uses
under outer, exfoliating layers. Leaves subopposite,
Large trees of this species are highly valuable tim- thick, multinerved, coriaceous, glabrous, (yellow-
ber trees, yielding large sizes of straight, knot-free, ish?) green. Leaves on saplings and young trees and/
strong and light coloured sawn timber. It is used or in shade ovate-lanceolate, up to 8 cm long and
for construction as beams, joists and frames, in car- 3 cm wide, acute or slightly acuminate; leaves in
pentry for joiner’s work, for boat building including crowns of mature trees distinctly petiolate, ovate to
oars due to its elasticity, idem for light aircraft as ovate-lanceolate, (2–)3–5(–7) cm long, (1–)2–3 cm
a substitute for Sitka spruce, and for floor boards, wide, with an obtuse or rounded apex. Pollen cones
panelling and furniture as well as picture frames, axillary, solitary on a (2–)5–10(–15) mm long, stout
152 pencils, rulers and T-squares. Lower grade wood is peduncle, when immature a small cylinder, 1–2 cm
used as pulpwood in the paper industry. In parts of long, 6–8 mm diam., elongating past anthesis to 3–4
Sulawesi where this tree no longer occurs subfossil cm long and up to 10 mm wide, with 4 decussate (in
resin (‘copal’) is found and mined, most likely pro- 2 pairs) 2–4 mm wide, free spreading bract scales
duced by trees that lived there many centuries ago. at their base, subtended by two much longer (up to
This hardened resin is the source for products like 30 × 10 mm) leaf-like bracts. Microsporophylls in
paints and varnishes and is searched for and dug imbricate arrangement; head convex towards the up-
up by ‘copal diggers’ as itinerant labourers setting per, rounded or retuse, minutely erose-denticulate
up camps in the forest, often following logging op- margin, 2 mm wide and 1.5 mm high in mature cones,
erations. This resin was formerly a much more im- bearing 3–6 oblong pollen sacs. Seed cones solitary on
portant product of this tree to the local inhabitants thick peduncles, broadly ellipsoid to globose, to 7–8
than its wood. Agathis dammara is used in forestry cm diam., with bossed scales (surface not smooth),
plantations, mainly in Jawa, where the genus does green, resinous, ripening brown. Cone scales with
not occur naturally. It is very rare in tropical botanic rounded to angular or bossed, thick upper margins,
gardens, where trees labeled A. dammara may be ca. 3 cm long, 4 cm wide in middle part of full-grown
A. borneensis instead. cones, rounded triangular to broadly reniform and
more or less flanged on either side. Seeds 11–13 × 7–8
mm, ovoid and flattened, with two unequal wings;
Agathis flavescens Ridl., Kew Bull. 1914: 332. 1914. largest wing ca. 15 × 10 mm; smallest wing a small
Agathis dammara (Lamb.) Rich. & A. Rich. subsp. blunt triangle 3 mm wide opposite largest wing.
flavescens (Ridl.) Whitmore, Pl. Syst. Evol. 135
(1–2): 59. 1980; Agathis celebica (Koord.) Warb.
Taxonomic notes
subsp. flavescens (Ridl.) Veldkamp & Whitmore,
Taxon 33 (2): 346. 1984. Type: Malaysia: Peninsular Whitmore (1980) recognized this taxon as a subspe-
Malaysia, Pahang, Gunung Tahan, H. N. Ridley cies of Agathis dammara, citing as the only mor-
16023 (holotype K). phological differences the two leaf-like bracts and a
slightly larger maximum size of pollen cones in sub-
species flavescens. However, the seed cone scales in
Etymology
A. flavescens often have an upper angular or bossed
The species epithet (Latin flavescens = yellowish or margin, giving the cone a more or less rough, not
pale yellow) refers to leaf colour. smooth surface, as in A. dammara. The leaves are,
though of course variable, smaller than in A. dam-
mara; but whether this is a trait, like the smaller size
Vernacular names
of trees, dependent on a more exposed environment
Tahan agathis. at high altitudes, can only be solved experimentally
by growing them under equal conditions. This has
not been done because A. flavescens, unlike A. dam-
Description
mara, is not of economic importance and is therefore
Trees to 18(–21) m tall and 1.5 m or more d.b.h. with not being grown in plantations. The often yellowish
short cylindrical bole and a spreading, rounded or colour of the leaves has been attributed to environ-
more or less flat-topped crown. Bark smooth or dip- mental conditions, e.g. nutrient deficiency in the
soil. If so, that would not be a taxonomic character,
Vernacular names
as it would change to green given access to more nu-
trients. Its isolated occurrence within the broader tumu (Sabah)
range of A. borneensis, not within that of A. dam-
mara, indicates that it will be found to be genetically
Description
isolated from both and is a distinct species.
Shrubs or trees to 20(–36) m tall and 1.5 m d.b.h.
Distribution with slender or short cylindrical bole and a spread-
ing, rounded or more or less flat-topped crown.
Malaysia: Peninsular Malaysia (Gunung Rabong
Bark smooth or dippled and lenticellate, colour
and Gunung Tahan).
grey, reddish brown under outer, exfoliating layers. 153
TDWG codes: 42 MLY-PM
Leaves subopposite, thick, multinerved, coriaceous,
glabrous, light green. Leaves on saplings and young
Ecology
trees and/or in shade ovate, up to 9 cm long and 4.5
Agathis flavescens occurs in the summit areas of two cm wide, (strongly) acuminate; leaves in crowns of
isolated mountains, at altitudes between 1100 m mature trees distinctly petiolate, ovate to obovate,
and 1900 m a.s.l. They are usually emergents above occasionally ovate-lanceolate, (2–)3–5(–7) cm long,
low mossy forest or occur as scattered small trees (1–)2–3.2 cm wide, with an obtuse or rounded, oc-
in mountain scrub. On Gunung Tahan, the highest casionally slightly acuminate apex. Pollen cones
mountain at 2189 m, they do not reach the summit, axillary, solitary on a 2–5 mm long, stout peduncle,
which is mostly covered in heath-like dwarf shrubs. when immature a small cylinder, 1–1.5 cm long, 6–8
Agathis borneensis occupies a lower zone on this mm diam., elongating past anthesis to 2.5–3 cm long
mountain covered with taller forest; the two appear and up to 10 mm wide, with 4–6 decussate (in 2–3
to be separated by a belt of vegetation devoid of Aga- pairs) 2–4 mm wide, free spreading bract scales at
this (Chung-Lu Lim, FRIM unpublished data). their base, subtended by two narrow, leaf-like bracts
to 20 × 4 mm. Microsporophylls in imbricate ar-
rangement; head slightly convex towards the upper,
Conservation
slightly angled, minutely erose-denticulate margin,
This species is restricted to isolated populations on 1.6–1.8 mm wide and 1.2–1.6 mm high in mature
two (possibly three) mountains. There may be fewer cones, bearing 3–6 oblong pollen sacs. Seed cones
than 10,000 mature trees existing in total. There is solitary on thick peduncles, broadly ellipsoid to sub-
no exploitation of this species for timber because the globose, to 11 cm long and 7–8.5 cm diam., smooth,
trees are small with mostly very short boles and they green, resinous, ripening brown. Cone scales with
are on road-less mountains. rounded, thick upper margins, ca. 3 cm long, 4–4.5
IUCN: VU (D2) cm wide in middle part of full-grown cones, round-
ed triangular and more or less flanged on either side.
Seeds 11–13 × 7–8 mm, ovoid and flattened, with two
Uses
unequal wings; largest wing 15–20 × 10–12 mm;
No uses have been recorded of this species. It is not smallest wing a small blunt triangle 3 mm wide op-
known to be in cultivation. posite largest wing.

Taxonomic notes
Agathis kinabaluensis de Laub., Blumea 25 (2): 535.
1979. Type: Malaysia: Sabah, Ranau District, Mt. Herbarium collections of this species from Mt.
Kinabalu NP., Summit Trail, D. J. de Laubenfels P Kinabalu go back at least to 1915; those from Gu-
625 (holotype L). Fig. 31, 32 nung Murud to 1970. De Laubenfels (1979, 1988)
­appears to have overlooked the latter collections in
his accounts of the genus in Borneo, while Whit-
Etymology
more (1980) seems to have included all of them
The species epithet refers to Mt. Kinabalu, from in Agathis dammara. The latter author did recog-
where it was first described. nize A. dammara subsp. flavescens as distinct from
 A. dammara subsp. dammara based on the elon- outside perhaps a few specimens in regional botanic
gated pair of bracts subtending the pollen cones, gardens.
but this character is also present in A. kinabaluensis
(they often have been broken off in dried herbarium
specimens). Agathis flavescens and A. kinabaluensis Agathis labillardierei Warb., Monsunia 1: 183. 1900,
are hardly different morphologically: the distinc- [“labillardieri”]. Type: Papua New Guinea, J. J. H. de
tions are minor and variable; perhaps with the ex- la Labillardière B-W 17793 (holotype B-W).
ception of acuminate leaves, which are not found in
A. flavescens, and of more or less bossed seed cone
Etymology
scales, not seen in A. kinabaluensis.
154 This species was named after Jacques Julien Houtton
de Labillardière (1775–1834), French explorer of the
Distribution
Pacific.
Borneo: Sabah (Mt. Kinabalu), Sarawak (Gunung
Murud).
Vernacular names
TDWG codes: 42 BOR-SB
New Guinea kauri; kayu damar putih (Indonesia,
general); kessi, fuko and many other local names,
Ecology
several cited in Flora Malesiana, ser. 1, 10 (3): 442
Agathis kinabaluensis is found in upper montane (1988) (New Guinea).
forest, mossy low forest and subalpine scrub, at alti-
tudes between 1500 m and 2400 m a.s.l. At the most
Description
exposed and highest sites it has a shrubby habit with
hardly any length of a single trunk, but in taller for- Trees to 60 m tall and 2 m or more d.b.h. with
est at lower altitudes it can become a tall tree itself. clear cylindrical bole the height of the canopy and
It occurs on nutrient-poor substrates such as ultra- a rounded or open crown emerging. Bark smooth,
mafics, granite or sandstone, in Fagaceae-Lauraceae exfoliating in irregular scales of up to 10 cm across;
dominated forest or stunted forest to scrub with outer bark brown, purplish under newly fallen scales;
Myrtaceae and numerous epiphytes. inner bark pinkish, exuding copious white resin.
Leaves subopposite, thin or thick, multinerved, co-
riaceous, glabrous, light green. Leaves on saplings
Conservation
and young trees and/or in shade broadly lanceolate,
This species is now recognized to occur on (at least) 9–14 cm long, 3–6 cm wide; leaves in crowns of ma-
two major mountains in N Borneo, but the popula- ture trees ovate-lanceolate or elliptic, 5–12 cm long,
tions are relatively small and could be negatively im- 1.5–3 cm wide, distinctly petiolate and with an acute
pacted by stochastic events such as forest fires. Some or obtuse apex; however some may be linear-lanceo-
subpopulations at lower altitude are under pressure late, 8–11 cm long and 1–1.5 cm wide. Pollen cones
from land-use conversion, in particular agriculture. axillary, solitary on a 2–6(–10) mm long, stout pe-
The population in Mt. Kinabalu National Park is duncle, short cylindrical or barrel-shaped, when full
under legal protection, but the other population on grown 2.2–3 cm long, 10–12 mm diam. with a round-
Gunung Murud is not. ed apex, with 8 decussate (in 4 pairs), imbricate but
IUCN: VU (D2) spreading, 4–6 mm wide bract scales at their base.
Microsporophylls in tesselate arrangement; head
rectangular to hexagonal, prismatic, with a raised,
Uses
flat central part 1–1.5 mm wide, 0.7–1 mm high in
No economic used have been reported of A. kina- mature cones; margins hidden. Seed cones solitary
baluensis, but where larger trees occur(ed) in the on thick peduncles, subglobose or obovoid, 8–10 cm
forest, they may have been logged by local people long, 7.5–9 cm wide, smooth, green and resinous,
to use the timber for construction and carpentry ripening brown. Cone scales with slightly rounded,
work. This species is not known to be in cultivation strongly incurved upper margins, ca. 3 cm long, 3.5–
4 cm wide, roughly triangular with rounded corners
and more or less flanged on either side. Seeds 12 × 7 banned in an attempt to provide work for local saw-
mm, ovoid, flattened, with two unequal wings; larg- mills and increase the value of timber. The wood is
est wing ca. 20 × 15 mm; smallest wing a small acute used for indoor construction, carpentry, boat build-
triangle 3–4 mm wide opposite largest wing. ing, veneer and tool making. In remote villages,
where modern sawing equipment is not available,
planks are sometimes split to build house walls. The
Distribution
resin (copal) is tapped and is an important compo-
New Guinea, including major islands around Vogel­ nent of varnish; more local and traditional uses in-
kop Peninsula and eastwards as far as the Sepik clude burning it as torches and for incense and other
­Valley. forms of divination.
TDWG codes: 43 NWG-IJ NWG-PN 155

Agathis lanceolata (Sébert & Pancher) Warb.,

Ecology
Monsunia 1: 186. 1900. Dammara lanceolata Sébert
Agathis labillardierei occurs in tropical evergreen & Pancher, Notes Bois Nouv. Calédonie: 169. 1874,
lowland to lower montane rainforest, in which it is non Vieill. (1862); Salisburyodendron lanceolata
an emergent tree. Locally it can be abundant in early (Warb.) A. V. Bobrov & Melikyan, Komarovia 4: 63.
phases of regeneration, but eventually angiosperms 2006. Type: New Caledonia: Grande Terre, Cougui
increase once more and close the canopy, leaving a (Mt. Koghis), J. A. I. Pancher s.n., 1870. (holotype
few solitary trees of A. labillardierei rising above it. P).
The altitudinal range of this species is from near sea
level to 1830(–2500?) m a.s.l., most trees occur be-
Etymology
tween 200 m and 1350 m a.s.l. This species is found
to grow on a variety of substrates, including ultra­ The species epithet comes from Latin lanceolata =
basic rock which produces soils poor in nutrients lanceolate; in reference to the shape of the leaves,
essential to tree growth. which (especially when young) are often in the
shape of a lance point.
Conservation
Vernacular names
This species has a wide distribution and is still com-
mon in many areas where the tropical rainforest of Koghis kauri, kaori
New Guinea remained intact to this day. It is to be
expected, however, that as similar resources else-
Description
where in Malesia are being exploited towards gen-
eral scarcity of large trees with long boles and with Trees to 40(–50) m tall and 1 m or more d.b.h. with
relatively easy access, the pressure on New Guinea clear cylindrical bole the height of the canopy and
kauri will increase. The ban on export of round logs a broad crown radiating above. Bark extremely
from Papua New Guinea will work in favour of more smooth, peeling in patches of up to 20 cm across
sustainable use, but most of the megapopulation of from the trunk, falling often and accumulating in
this species occurs in Papua (the Indonesian half of large piles around the base of the tree; colour vari-
New Guinea), where this ban has not (yet) been im- able – in forest deep red, but becoming greyer (while
posed. It is perhaps prudent to upgrade the conser- ­remaining smooth) with exposure to sun. Leaves
vation status of this species in anticipation of these subopposite, thick, multinerved, coriaceous, gla-
developments. brous, light green. Leaves on saplings and young
IUCN: LC trees and/or in shade lanceolate to falcate, 9–13 cm
long, 3–5 cm wide; leaves in crowns of mature trees
ovate-lanceolate, 6–8 cm long, 1.5–2 cm wide with
Uses
an obtuse apex; however some may be long and
New Guinea kauri is one of the most valuable timber linear, ca. 8–10 cm long and 1.5 cm wide. Pollen
trees of the island. While logging continues, export cones axillary, solitary on a 4–5 mm long, stout pe-
of round logs from Papua New Guinea has been duncle, cylindrical or nearly oval, when full grown
 2–2.5 cm long, 8–10 mm diam., with 8 decussate (in
Uses
4 pairs), free, 4 mm wide bract scales at their base,
the lowest pair often developed into small leaflets. Timber of this tree is extremely valuable: the species
Micro­sporophylls in tesselate arrangement; head is widely used in reafforestation projects in southern
prismatic, forming a bossed surface, less than 1.6 New Caledonia. It was logged heavily in the past and
mm wide, 0.5–1.75 mm high in mature cones; mar- used for carpentry and boat building; as large trees
gin entire. Seed cones solitary on thick peduncles, in the wild are no longer felled, the smaller sizes
globose, 10–12 cm diam., smooth, green or glaucous of planted trees have to be used instead. Resin was
green, (very) resinous, ripening brown. Cone scales ­another major product from the old stands, largely
with slightly rounded, incurved upper margins, ca. going to the turpentine and varnish industries.
156 2.5 cm long, 3–3.5 cm wide, roughly triangular with
rounded corners and more or less flanged on either
side. Seeds 12–15 × 7 mm, ovoid-oblong, with two Agathis lenticula de Laub., Blumea 25 (2): 537.
unequal wings; largest wing ca. 20 × 13 mm; smallest 1979. Type: Malaysia: Sabah, Ranau District, Mt.
wing a small blunt triangle 3–5 mm wide opposite Kinabalu N. P., near Park Headquarters, D. J. de
largest wing. Laubenfels P 619 (holotype L). Fig. 33, 34

Distribution Etymology
New Caledonia: Grande Terre, mainly in Province The species epithet (Latin lenticularis = lens-shaped)
Sud (southern massif). refers to the leaves.
TDWG codes: 60 NWC
Vernacular names
Ecology
tangilan (and variant spellings) (Sabah).
Agathis lanceolata occurs mostly in rain forest, often
in valley bottoms and can occasionally form rela-
Description
tively pure stands there, or it grows together with
Araucaria subulata; sometimes it is associated with Trees to 40(–45) m tall and 2.2 m or more d.b.h., with
Nothofagus spp. This species is also found, possibly clear cylindrical bole up to 20–25 m and a broad crown
in relict stands following 19th century logging, in radiating above. Bark variable; usually lenticellate
isolated pockets of humid forest (often with Aril- and scaly with large but irregular flakes, colour grey
lastrum gummiferum, Montrouziera cauliflora, and to dark brown or greenish purple outside, reddish
Calophyllum spp., or sometimes with Podocarpus brown or yellowish brown under outer, exfoliating
sylvestris) in the hollows of valley heads of the south- layers, exuding clear or white resin turning yellow.
ern massif around the Plaine des Lacs, an area oth- Leaves subopposite, thick, multinerved, coriaceous,
erwise dominated by maquis, and in areas where the glabrous, light green and often glaucous underneath.
boundary between forest and maquis is not sharp. It Leaves on saplings and young trees and/or in shade
is a species of lower altitudes, between 50 m and 500 broadly lanceolate to elliptic, 6–11 cm long, 2–4.5 cm
m a.s.l., rarely up to 900 m a.s.l. It usually grows on wide, acute or slightly acuminate; leaves in crowns of
red ultramafic soils. mature trees distinctly petiolate, lenticular or ovate-
elliptic, 5–8 cm long, 1–2.5 cm wide, acute or obtuse.
Pollen cones axillary, solitary on a 2–10 mm long,
Conservation
stout or slender peduncle, when immature from sub-
Historically, this species has been seriously over­ globose becoming nearly cylindrical, 1–1.5 cm long,
exploited for both timber and resin. Some popu- 7–8 mm diam., elongating past anthesis to 3–4 cm
lations are now protected in several reserves, long and 9–10 mm wide, with 4–6 decussate (in 2–3
including major reserves, across the southern part pairs) 3–4 mm wide, imbricate bract scales at their
of the country, but ongoing decline is projected to base. Microsporophylls in imbricate arrangement;
occur elsewhere. head slightly convex towards the upper, rounded to
IUCN: VU [B1ab(iii)+2ab(iii), C2a(i)] angular, minutely erose-­denticulate margin, 2–2.5
mm wide and 1.5–2 mm high in mature cones, bear- boles, must be similar outside protected areas. If we
ing 3–6 oblong pollen sacs. Seed cones solitary on would know more about specific rates of exploita-
thick peduncles, globose, to 8–10 cm diam., more or tion in particular areas where A. lenticula has been
less rough with raised seed scale margins, especially found, it could well be that A. lenticula turns out to
in immature cones, green, resinous, ripening brown. be more seriously threatened than the more wide-
Cone scales with more or less rounded, bossed, thick spread species A. borneensis.
and incurved upper margins, 2.5–3 cm long, 3–3.5 IUCN: VU (D2)
cm wide in mature cones, roughly flabellate with
rounded corners and more or less flanged on either
Uses
side. Seeds ca. 10 × 6 mm, ovoid, flattened, with two
unequal wings; largest wing ca. 15 × 8 mm; smallest This species is not distinguished from A. borneen- 157
wing a small, blunt triangle of 3 mm opposite large sis by foresters and will be logged as that species or
wing, or absent. ‘dammar’ or ‘kauri’ timber. Its uses are similar to
those of A. borneensis.
Taxonomic notes
Whitmore (1980) included herbarium collections of Agathis macrophylla (Lindl.) Mast., J. Roy. Hort.
this species in a broader concept of Agathis damma- Soc. London 14: 197. 1892. Dammara macrophylla
ra, which although having a distribution generally Lindl., J. Hort. Soc. London 6: 271. 1851. Type:
to the east of A. borneensis, in his opinion occupied Solomon Islands: Santa Cruz Group, Vanikoro
several isolated montane stations within the range Island, Vanikolo, [“Hab. Island of Vanikolla”],
of A. borneensis. These isolated populations were C. Moore s.n. (holotype CGE).
all recognized as distinct species by De Laubenfels
(1979, 1988) in a nearly simultaneous taxonomic Dammara obtusa Lindl., J. Hort. Soc. London 6: 270.
study of the genus in Malesia. 1851; Agathis obtusa (Lindl.) Mast., J. Roy. Hort. Soc.
London 14: 197. 1892; Agathis macrophylla (Lindl.)
Mast. var. obtusa (Lindl.) Silba, Phytologia 68: 23.
Distribution
1990.
Malaysia: Sabah (Crocker Range to Mt. Kinabalu). Dammara brownii hort. ex Lem., Ill. Hort. 2, Misc.:
TDWG codes: 42 BOR-SB 60. 1855; Agathis brownii (Lem.) L. H. Bailey, Cult.
Conif. N. Amer.: 18 passim, 151. 1933.
Dammara vitiensis Seem., Fl. Vitiensis: 265, t. 76.
Ecology
1868; Agathis vitiensis (Seem.) Benth. & Hook. f.,
Agathis lenticula is an emergent tree in lower mon- Gen. Pl. 3 (1): 436. 1880.
tane evergreen tropical rainforest. It occurs in dipte-
rocarp rainforest and forest dominated by Fagaceae
Etymology
at altitudes between 1050 m and 1700 m a.s.l. on a
variety of dark to light soils. The species epithet comes from Greek macrophylla
= ‘with large leaves’; compared with most other Aga-
this species, this species has especially large leaves.
Conservation
This species has been assessed as Vulnerable pri-
Vernacular names
marily on the basis of a limited distribution (area
of occupancy less than 100 km² under IUCN 1994 Fijian kauri pine; dakua, ndakua, ndakua makandre
criteria) because it is very difficult to estimate its (Fiji); nendö, notopiti (Santa Cruz Group); kauri
decline due to exploitation. Most herbarium collec- (Vanuatu).
tions at K from the Crocker Range and lower spurs
and slopes of Mt. Kinabalu in Sabah have been iden-
Description
tified as A. lenticula, but A. borneensis does occur
in the region and foresters do not distinguish them. Trees to 35(–40) m tall and 1 m or more d.b.h. with
Exploitation of both, being tall trees with long, free clear but usually short bole and an ultimately broad
 crown formed by long, spreading and ascending phylla, for these entities. This excludes trees more
branches. Bark very scaly, exfoliating in irregu- recently discovered on Santo Peak, Espiritu Santo,
lar patches of variable shape and size; colour grey. in Vanuatu, which are a distinct species A. silbae.
Leaves subopposite, thick, multinerved, coriaceous, Such ­discoveries may hint at the possibility of one or
glabrous, light green or glaucous to farinose esp. more other ‘hidden’ distinct species on these islands,
on the abaxial (lower) side. Leaves on saplings and but such have not been observed among the mate-
young trees and/or in shade broadly lanceolate, rial available to researchers at this time.
sometimes slightly falcate, 8–17 cm long, 3–6 cm
wide, with acute or obtuse apex; leaves in crowns
Distribution
of mature trees ovate-lanceolate to obovate or occa-
158 sionally nearly round, (2.5–)4–8 cm long, (0.8–)1.5–3 Fiji: Kadavu Is., Viti Levu, Vanua Levu; Solomon Is-
cm wide with an obtuse or round apex. Pollen cones lands: Santa Cruz Group (Utupua Is., Vanikoro Is.);
axillary, solitary on a (0–)3–7 mm long, stout pe- Vanuatu: Anatom Is., Erromango Is., Tanna Is.
duncle, mostly becoming cylindrical, (2–)2.5–4.5 cm TDWG codes: FIJ SCZ VAN
long, 8–15 mm diam., with 8 decussate (in 4 pairs),
imbricate, 4–7 mm wide bract scales at their base,
Ecology
spreading to form a basal collar somewhat wider
than the pollen cone base. Microsporophylls in im- Agathis macrophylla is an emergent tree in lowland
bricate arrangement; head convex, 1.4–2.2 mm wide, to low montane tropical rainforest; usually growing
1.8–2.2 mm high in mature cones; margin minutely in soils derived from volcanic rocks like basalt. Its
erose-denticulate, often notched at apex. Seed cones altitudinal range is recorded from herbarium col-
solitary on thick peduncles, globose, 10–13 cm diam., lections as being between 75 m and 900 m a.s.l. In
smooth, green or glaucous green, (very) resinous, Fiji on the main islands it is most common between
ripening brown. Cone scales with slightly rounded, 600 m and 900 m a.s.l. The species was the subject
incurved upper margins, ca. 3.5 cm long, 3.5–4.5 cm of a pioneering study on its role in the rain forests
wide, roughly triangular with rounded corners and of the type locality, which demonstrated that unlike
more or less flanged on either side. Seeds 12–15 × 7–8 Agathis australis in New Zealand, and many other
mm, ovoid-oblong, with two unequal wings; largest species which exhibit a regeneration strategy based
wing 20–25 × 10–15 mm; smallest wing a small blunt on periodic landscape-scale disturbance (cf. Enright
triangle 3–6 mm wide opposite largest wing. & Hill, 1995) A. macrophylla appears to behave as a
normal component of rainforests dominated by an-
giosperms. This means that it is capable of small-gap
Taxonomic notes
regeneration like other large forest trees.
Foresters and others in the SW Pacific familiar with
this tree tend to recognize two or more distinct en-
Conservation
tities (species), especially a distinction between the
trees in Fiji and those in the other island groups. Overall, this species is listed as Near Threatened
Taxonomists (e.g. Whitmore, 1980) are more im- (IUCN Redlist). However, individual island popula-
pressed by the continuous variation observed in tions may well be severely threatened, e.g. that on
numerous herbarium specimens collected from all Utupua in the Santa Cruz Group. Doyle (in Farjon &
islands where Agathis occurs. There is perhaps no Page, 1999) assessed the species separately for each
other species in this genus with more variable foli- of the island groups as follows: Fiji: VU – Vulnera-
age leaves than A. macrophylla; especially expressed ble, IUCN 1994-criteria A2d (>20% reduction within
in canopy foliage. Specimens from Fiji are often next three generations based on current exploitation
glaucous to farinose on one side of the leaves, but levels) and B2e (continuing decline in the number
non-glaucous specimens have been collected on of mature individuals). Santa Cruz Islands: NE –
these islands, too. Characters that really matter in not evaluated. Vanuatu: VU – Vulnerable, IUCN
this difficult genus, like those of the pollen cones, are 1994-criteria A2d and B2e. Agathis macrophylla is
quite uniform across the island groups. I therefore now quite rare outside plantations in the Santa Cruz
concur with Whitmore (1980) and here recognize Group, though an unlogged population apparently
one species, under its earliest name, A. macro- survives on the upper Lawrence River on Vanikoro.
It seems likely that a re-assessment using version 3.1 crown with spreading to ascending branches. Bark
of the IUCN criteria (IUCN, 2000) would conclude smooth or sometimes scaly, exfoliating in irregu-
on a threatened category, probably VU, for the entire lar coarse scales of up to 15 cm across; outer bark
known population. brown or grey-brown, with bluish purple patches
IUCN: NT under newly fallen scales; inner bark reddish, exud-
ing white resin. Leaves subopposite, thin or thick,
multinerved, coriaceous, glabrous, light green.
Uses
Leaves on saplings and young trees and/or in shade
The wood of this species is white or sometimes with broadly lanceolate to elliptic, 5–9 cm long, 1.5–3.5
a reddish hue and known in Fiji as Dakua wood and cm wide; leaves in crowns of mature trees lanceolate
in the Santa Cruz Group as Vanikoro kauri. It is very to elliptic, 3–8 cm long, (0.8–)1–2.2 cm wide, short 159
valuable and used for construction, for flooring in petiolate and with an obtuse or rounded apex. Pol-
houses, for masts, booms and spars in sailing boats, len cones axillary, solitary on a 0–1 mm long pedun-
for carpentry and for furniture making. The resin cle, subglobose becoming short cylindrical, when
exuded from the bark is fragrant and inflammable full grown 1–1.6 cm long, 5–8 mm diam., with 8–10
and is (was) burnt to provide light. Recent resin is decussate (in 4–5 pairs), imbricate, 1–2.5 mm wide
transparent and nearly colourless but weathers white bract scales at their base. Microsporophylls in tes-
in contact with air and sunlight; subfossil resin has selate arrangement; heads prismatic, with raised
a yellowish or orange-brown hue approaching some central part irregularly pentagonal or hexagonal and
types of amber, which is completely fossilized (‘ma- becoming free at anthesis, 0.5 mm wide, 0.5–0.7 mm
tured’) resin from conifers. Resin is tapped from high in mature cones, bearing 2–5 pollen sacs. Seed
trees, but also dug from the ground (subfossil resin) cones solitary on thick peduncles, globose or ovoid,
and used in making varnishes, pottery glazing, and 7.5–11.5 cm long, 6.5–10 cm wide, smooth or nearly
dying cloth black with the smoke from burning it. so, green or slightly glaucous, ripening brown. Cone
Fijian kauri pine has been planted as a forestry tree scales with thick, rounded, incurved upper margins,
in the Solomon Islands (Santa Cruz Group) and 2.5–3.5 cm long, 3.3–4.5 cm wide, broad triangular
elsewhere in the SW Pacific in an attempt to obtain with rounded corners and more or less flanged on
timber more sustainably from a truly renewable re- either side. Seeds 10–12 × 6–8 mm, ovoid, flattened,
source. It is also in cultivation in some greenhouses with two unequal wings; largest wing 20–25 × 10–13
of botanic gardens. mm; smallest wing a more or less triangular point
2–5 mm long, opposite largest wing.
Agathis microstachya J. F. Bailey & C. T. White,
Contr. Queensland Fl. Bot. Bull. 18: 13. 1916. Taxonomic notes
Type: Australia: Queensland, Cook District,
Agathis atropurpurea and A. microstachya are more
H. W. Mocatta s.n. (holotype BRI). Fig. 35
or less sympatric, but remain often separated from
each other by altitudinal range (with A. micro-
Etymology
stachya usually occurring at a lower altitude), and
The species epithet derives from the Greek micros = slight habitat differences, e.g. soil conditions, which
small and stachys = ear of corn (maize, not known in are generally poorer with A. atropurpurea. They
ancient Greece) or a flower spike, and alludes to the do not seem to hybridize. The most consistent dif-
small male strobili. ference between them is the morphology of the
micro­sporophylls in the pollen cones (these cones
are small in both species). In A. atropurpurea, these
Vernacular names
are imbricately arranged (the most common situ-
Bull kauri, Bull pine, Atherton kauri pine ation in the genus), while the microsporophylls in
A. microstachya are tesselate, i.e. laying together like
paving stones, with no parts overlapping. The shapes
Description
of the microsporophyll heads are also markedly dif-
Trees to 50 m tall and 2.5 m or more d.b.h. with ferent. The seed cones of A. microstachya are, when
clear cylindrical bole to 35 m and a rounded or open mature, much larger than the smallish seed cones
 of A. ­atropurpurea. There is a general leaf shape dif- Agathis montana de Laub., Trav. Lab. Forest. Tou­
ference in leaves on mature trees in that A. micro- louse T. 1 (8, 5): 2. 1969. Salisburyodendron montana
stachya has obtuse or rounded leaf apices, while (de Laub.) A. V. Bobrov & Melikyan, Komarovia
A. atropurpurea has only obtuse apices, but this 4: 63. 2006. Type: New Caledonia: Grande Terre,
character is less reliable. Province Nord, Mt. Panié, M. Schmid 1420 (holo­
type P).
Distribution
Etymology
Australia: NE Queensland (Cook District).
TDWG codes: 50 QLD-QU The species epithet means “from the mountain” and
160 refers to its habitat.
Ecology
Vernacular names
Agathis microstachya occurs mainly scattered in
lowland to low montane semi-evergreen tropical No common names are known for this species.
rainforest in mountains E of the Atherton Table-
lands, but also less frequently on these plateaus. Its
Description
altitudinal range is between 400 m and 1100 m a.s.l.
It is an emergent species in a species-rich canopy Medium size trees to 15–20 m tall, either multi-
of angiosperm trees. Soils are loamy sands or light stemmed from near base or with clean bole for 8–10
clays and derived from acidic to neutral silicate-rich m; sometimes almost flat-topped in appearance but
rock, usually of volcanic origin. Annual rainfall var- crown usually very sparse, with long, spreading,
ies, with maxima to over 3000 mm on ocean-facing crooked branches. Young trees possibly conform-
mountain slopes. ing to Rauh’s architectural model, with whorled
tiers of assurgent branches forming a conical crown.
Bark reddish brown or tan to grey with numerous
Conservation
small, granular lenticels, flaking, coming away from
Before 1985 the population of A. microstachya had the tree in irregular small pieces, excuding copious
been halved by logging but 70% of the forests are white resin. Leaves subopposite, thick, multi­nerved,
now protected. The remaining population is esti- coriaceous, glabrous, glaucous on the abaxial sur-
mated to consist of fewer than 10,000 mature trees. face. Leaves of saplings and young trees broadly
Under IUCN Red List criteria (version 3.1, 2001) ovate, 8–11 cm long, 2.4–3.8 cm wide; leaves in crown
this species would therefore qualify for the status of mature trees becoming more lenticular, 5.5–9 cm
VU, but due to cessation of large-scale logging and long, 1.4–2.2 cm wide. Pollen cones axillary, solitary,
effective protection in a National Park and other short pedunculate or sessile, with a tight basal-bract
protected areas, its threat with extinction has been cluster of 5–8 imbricate, 1–2 mm wide bract pairs and
greatly reduced. Logging continues in unprotected without a pair of leaf-like bracts, cylindrical, 4–5 cm
areas, but the decline appears to have been halted. long but expanding to 7 cm or more past anthesis,
IUCN: NT 8–10 mm wide. Microsporophylls imbricate; micro-
sporophyll heads umbonate, apices weakly acumi-
nate, 2.0–2.6 mm wide and 2.2–2.5 mm high. Seed
Uses
cones axillary, solitary on stout peduncle, globose to
Bull kauri is a valuable timber tree and was formerly obovoid, ca. 9 cm long, 7 cm wide, smooth, green to
intensively logged and taken to local sawmills. The glaucous green ripening brown, very resinous. Cone
timber is soft, light, easy to work and polishes well. scales with rounded, thick and slightly incurved
It is used for house framing, flooring, and joinery, upper margins and thin, fragile lateral margins, ca.
as well as veneer production. This species is planted 3 cm long and wide. Seeds ovoid, 6–8 × 5 mm, with
on a limited scale mainly in arboreta and tree collec- one large ovate wing and a small, triangular wing on
tions in parks in Queensland. the opposite side (no intact wings seen).
 Distribution Vernacular names
New Caledonia (Mt. Panié). Moore’s kauri; kaori blanc, kaori rouge (French in
TDWG codes: 60 NWC New Caledonia)

Ecology Description
Agathis montana is abundant to dominant from Trees to 40(–50) m tall and 1 m or more d.b.h. with
1100–1200 m upwards on the Panié massif, the clear cylindrical bole to 25 m; crown conical or
highest and wettest mountains in New Caledonia. rounded in trees of medium size, becoming more
It occurs in fern-rich rain forest habitat with Retro­ open and irregular in old trees. Bark smooth or 161
phyllum comptonii, the only other large tree, on rough and scaly, peeling in small or large, irregu-
reddish soil overlying micaschists. On the summit lar flakes; colour variable – in forest reddish brown
of Mt. Panié Agathis montana occurs together with to tan, but becoming greyer (and harder) with ex-
Araucaria schmidii on or near exposed outcrops of posure to sun, often resinous. Branches spreading.
rock and steep summit slopes, where it becomes Leaves subopposite, thick, multinerved, coriaceous,
stunted. The altitudinal range of this species is from glabrous, light green, sometimes glaucous on the
950 m a.s.l. to 1600 m a.s.l. on the summit crest. abaxial (lower) side. Leaves on saplings and young
trees and/or in shade lanceolate to elliptic, 10–20 cm
Conservation long, 2–4 cm wide; leaves in crowns of mature trees
narrowly ovate-elliptic, 4.5–7 cm long, (6–)8–12
IUCN: NT
mm wide with an obtuse apex. Pollen cones axil-
lary, solitary on a slender, (1–)8–12 mm long pedun-
Uses
cle, cylindrical, when full grown 2.5–3(–5) cm long,
No uses are recorded of this species. The first bo- 6–9 mm diam., with 8–16 decussate (in 4–8 pairs),
tanical collection is from as recently as 1939 and the imbricate, 4–5 mm wide bract scales at their base.
altitude and relative inaccessibility of all the known Microsporophylls in imbricate arrangement; head
stands may have prevented earlier utilization, e.g. more or less angular but nearly flat, 1–3 mm wide,
the collecting of resin. The commonly crooked 0.75–2 mm high in mature cones; margin minutely
shape and shortness of the boles render it less suit- erose-denticulate. Seed cones solitary on thick pe-
able for timber than most other species in the genus. duncles, subglobose to globose, 10–15 cm long, 9–12
It is not known to be in cultivation. cm diam., smooth, green or glaucous green, (very)
resinous, ripening brown. Cone scales with slightly
rounded, incurved upper margins, 3–4 cm long, 3–4
Agathis moorei (Lindl.) Mast., J. Roy. Hort. Soc.
cm wide, roughly triangular with rounded corners
London 14: 197. 1892. Dammara moorei Lindl.,
and more or less flanged on either side. Seeds 15–20
J. Hort. Soc. London 6: 271. 1851; Salisburyodendron
× 8–10 mm, ovoid-oblong, with two unequal wings;
moorei (Lindl.) A. V. Bobrov & Melikyan, Koma­
largest wing 20–30 × 15–20 mm; smallest wing re-
rovia 4: 63. 2006. Type: New Caledonia: Grande
duced to a narrow appendix 3–5 mm long, opposite
Terre, [“on East Coast”], C. Moore s.n. (holotype
large wing.
CGE).
Agathis corbassonii de Laub., Trav. Lab. Forest. Tou-
Taxonomic notes
louse T. 1 (8, 5): 2. 1969. Salisburyodendron corbasso-
nii (de Laub.) A. V. Bobrov & Melikyan, Komarovia Agathis corbassonii was segregated from A. moorei
4: 63. 2006. on the basis of its very reddish brown bark and its
narrow leaves which are often glaucous beneath.
The possibility of separate species (kaori blanc, ka-
Etymology
ori rouge) was first suggested by Michel Corbasson,
This species was named after Charles Moore (1820– Director of the Tropical Forestry Institute in Nou-
1905), a Director of the Royal Botanic Gardens Syd- mea. However, the variation in leaf sizes and shapes
ney, who collected the type specimen. in A. moorei is perhaps greater than in any other
 species in the SW Pacific (T. G. Waters, unpublished because illegal logging is a particular concern (Watt
D. Phil thesis, Oxford 2008) and those specimens as- in Farjon & Page, 1999). However, the species still
cribed to A. corbassonii appear to fall mostly within occurs over much of the island of Grande Terre and,
the ranges observed in A. moorei. Glaucousness is given adequate protection, could recover in many
not a sound character for taxonomic distinction as it areas to former abundance.
so often varies within populations in conifers. Field IUCN: VU [B1ab(iii)+2ab(iii). C1]
observation strongly indicates that shaded boles
within forests have reddish brown bark, while sun-
Uses
exposed trees develop grey bark through a weather-
ing process. This exposed bark also becomes harder Moore’s kauri is a valuable timber tree and its wood
162 than on trees in moist forests. [It is understandable is (was) much used for carpentry, joinery, veneer,
that local foresters took note of these bark differ- and for the construction of ‘pirogues’ (outrigger ca-
ences, which is often all one can see of a large tree noes). In the extreme southern part of Grande Terre
in tropical rainforest.] The number of bract scales some limited forestry plantations of this species
at the base of the pollen cones in A. moorei is not have been established on ultramafic soils.
consistently 16 (8 pairs in decussate arrangement) as
is stated in some published descriptions, but can be
fewer; it is a maximum number distinguishing the Agathis orbicula de Laub., Blumea 25 (2): 540.
pollen cones of A. moorei from other species in New 1979. Type: Malaysia: Sarawak, 5th Division, Lawas,
Caledonia. Agathis corbassonii is here treated as a [“Bumbong Rumah, N of Lawas”], D. J. Laubenfels
synonym of A. moorei. The lower number of bract de P 614 (holotype L).
scales recorded for A. corbassonii (based on few
good specimens) is therefore included in the char-
Etymology
acter states as found in A. moorei, and in agreement
with the measurements compiled by Waters and his The species epithet (Latin orbis = globe) describes
judgement on the matter. the shape of (some of) the leaves as being nearly
round.
Distribution
Vernacular names
New Caledonia: Grande Terre.
TDWG codes: 60 NWC tumuh (Murut); tubu (Kenyah); bulok (Iban)

Trees to 40 m tall and 2 m or more d.b.h., with clear

Ecology
cylindrical bole up to 20 m and a broad crown ra-
Agathis moorei occurs in dense rain forests on soil diating above. Bark sparsely lenticellate, hard and
derived from schists, micaschists, gneiss, and ser- smooth, exfoliating in small plates to 4 mm thick,
pentines. It is usually restriced to non-ultramafic colour dark brown or greenish purple outside, red-
soils, either forming small groves of pure Agathis dish brown under outer, exfoliating layers, exud-
moorei, or scattered through mixed angiosperm for- ing light yellow resin. Leaves subopposite, thick,
est. Its altitudinal range is from (30) 100 m to 700 multinerved, coriaceous, glabrous, light green and
(1000) m a.s.l. often glaucous underneath. Leaves on saplings and
young trees and/or in shade broadly lanceolate, 4–7
cm long, 2–3.5 cm wide, acute or slightly acuminate;
Conservation
leaves in crowns of mature trees distinctly petiolate,
This species has been heavily exploited for its tim- oval or ovate to nearly orbiculate, (2–)2.5–4(–6) cm
ber in the past and consequently it has disappeared long, 1–2.5 cm wide, acutish or obtuse to rounded
or become very scarce in many parts of the for- at apex. Pollen cones axillary, solitary on a 2–3 mm
est, especially in more accessible localities. Even in long peduncle, when immature from subglobose be-
pre-European times its wood was preferred for the coming nearly cylindrical, 6–8 mm long, 4–5 mm
making of outrigger canoes. Substantial decline has diam., only slightly elongating past anthesis to 12–15
occurred also within recent years and may continue mm long and 5–6 mm wide, with 4–6 ­decussate
(in 2–3 pairs) 2–3 mm wide, free spreading bract Borneo in Kalimantan Timur. Exploitation of all tall
scales at their base. Microsporophylls in imbricate trees of Agathis in Borneo has been intensive and is
arrangement; head slightly convex towards the up- still ongoing. As a consequence, we can infer a de-
per, rounded to angular, minutely erose-denticulate cline even though this species is not distinguished
margin, 1.2–1.5 mm wide and 1 mm high in mature by foresters or loggers from the more widespread
cones, bearing 2–4 oblong pollen sacs. Seed cones species A. borneensis, which occurs in the same gen-
solitary on thick peduncles, ovoid to subglobose, eral area. The true rate of decline could be impos-
to 8 cm long and 6 cm diam., more or less rough sible to estimate, as logged trees and their stumps
with raised seed scale margins especially in imma- are no longer identifiable to species and cut trees as-
ture cones, green, resinous, ripening brown. Cone sumed to belong to the more common A. borneensis
scales with more or less rounded, bossed, thick and may have been A. orbicula. It may have been present 163
strongly incurved upper margins, ca. 2 cm long and in several more than the presently known localities.
3 cm wide in mature cones, flabellate to nearly reni- IUCN: VU (B1+2c)
form with rounded corners and more or less flanged
on either side. Seeds ca. 10 × 6 mm, ovoid, flattened,
Uses
with two unequal wings; largest wing ca. 12 × 6 mm;
smallest wing a small triangle of 2 mm opposite The wood of this species has the same properties
large wing, or absent. as other lowland species of Agathis in Borneo and
is used for the same purposes; see therefore under
A. borneensis. The resin is said to have a yellow col-
Taxonomic notes
our, but since the resin of A. lenticula and other
Agathis orbicula has similar shapes and sizes of species comes out clear or white and tends to turn
leaves to A. kinabaluensis, but those of the latter are yellowish on exposure to air, this may be a some-
not glaucous on the abaxial (lower) side. The pol- what doubtful property and is not known to be rel-
len cones of A. orbicula are among the smallest in evant to its applications. This species is not known
the genus (after anthesis) and lack the two elongated to be in cultivation, but since plantations for forestry
basal bracts observed in A. kinabaluensis. Agathis have not made the taxonomic distinction between
orbicula occurs at lower altitudes in tall forest and this species and A. borneensis, it may be present in
consequently is a tall tree; A. kinabaluensis occurs at some plantations.
high montane altitudes and is mostly a short, stunt-
ed tree, but grows tall when in high forest.
Agathis ovata (C. Moore ex Vieill.) Warb., Mon­
sunia 1: 186. 1900. Dammara ovata C. Moore ex
Distribution
Vieill., Ann. Sci. Nat. Bot., sér. 4, 16: 56. 1862;
Borneo: Indonesia (Kalimantan Timur); Malaysia Salisburyodendron ovata (Vieill.) A. V. Bobrov
(Sabah, Sarawak). & Melikyan, Komarovia 4: 63. 2006. Type: New
TDWG codes: 42 Bor-ka bOR-SB BOR-SR Caledonia: Grande Terre, Province Sud, Col de
Yaté, [“Monts d’Unia”], E. Vieillard 1263 (holotype
P). Fig. 36, 37
Ecology
Agathis orbicula occurs at lower altitudes than Dammara hypoleuca C. Moore ex Henkel &
A. lenticula in evergreen tropical rainforest and high W. Hochst., Syn. Nadelhölz.: 217. 1865; Agathis hypo­
kerangas. It is, like that and other species at these leuca (C. Moore ex Henkel & W. Hochst.) Warb.,
lower altitudes (for this species between 450 m and Monsunia 1: 186. 1900; Salisburyodendron ovata
1050 m a.s.l.) a tall emergent tree. (Vieill.) A. V. Bobrov & Melikyan subsp. hypoleuca
(C. Moore ex Henkel & W. Hochst.) A. V. Bobrov &
Melikyan, Komarovia 4: 63. 2006.
Conservation
This species is known from six localities. In addi-
Etymology
tion to those mapped in Flora Malesiana, ser. 1, 10
(3): 437, f. 78 (De Laubenfels, 1988) there is one on The species epithet refers to the often ovate leaves.
 Vernacular names Distribution
Scrub kauri; kaori de montagne, kaori nain (French New Caledonia: Grande Terre, mainly in Province
in New Caledonia) Sud (Massif du Sud) but with two localities further
north, one as far as Commune de Houaïlou in Prov-
ince Nord.
Description
TDWG codes: 60 NWC
Small, often shrubby trees up to 10 m tall (occasion-
ally to 25 m in forest), usually with a short trunk
Ecology
to 80 cm d.b.h. and branching from near the base,
164 with orthotropic primary branches (Rauh’s model) Agathis ovata is usually growing on ultramafic soils
forming a flat-topped or irregular, spreading crown. in both maquis and (more rarely) forest environ-
Leaf-bearing branches and cones held in dense pseu- ments. Its altitudinal range is from 30 m to 1050 m
do-verticillate, upright or ascending clusters at tips a.s.l. In maquis environments it commonly occurs
of main branches, often glaucous to farinose. Bark with Araucaria spp., Babingtonia leratii, Xanthoste-
with deep fissures, hard, slowly flaking off and form- mon spp., Grevillea spp., various members of the Cy-
ing irregular polygonal patches and scales on main peraceae, Pteridium esculentum, Dracophyllum spp.,
trunks and branches, tan or reddish soon becom- various members of the Cunoniaceae, Myodocarpus
ing grey. Leaves subopposite, thick, multinerved, spp., Dacrydium araucarioides, Lomandra insularis,
coriaceous, glabrous, light green or glaucous green Gymnostoma spp., Styphelia spp., Asplenium spp.,
on the abaxial (under) side. Leaves on saplings or Hibbertia spp., and lichens such as Cladia retipora.
young trees well spaced, spreading in a plane, obo- In forest environments it occurs in mixed tropi-
vate-truncate, 4–6 cm long, 10–15 mm wide. Leaves cal rain forest, usually on drier, exposed slopes or
in crowns of mature trees crowded, often in 4 more ridges, but including forests otherwise dominated
or less erect rows, similar in shape, often somewhat by species of Nothofagus. Here it may occasionally
wider and with the leaf apex truncate or notched, attain 20–25 m and, due to competition with other
4–8.5 cm long and 1.5–5 cm wide but occasionally trees, grow as an upright tree.
almost orbiculate in shape. Pollen cones axillary,
solitary on a stout 4–20 mm long peduncle, more or
Conservation
less fusiform (widest in the middle) but with an ob-
tuse apex, 3–5 cm long, 10–15 mm diam.; subtended This species is widespread but scattered in ultrama-
by two small leaf-like bracts and 4–5 pairs of imbri- fic maquis in Province Sud, and protected in some
cate, short, rounded bract scales; basal-bract cluster major reserves. Fire remains a threat in many areas
usually narrower than widest point of pollen cone. where its incidence has been increased by humans,
Microsporophylls imbricate, microsporophyll head and a continuing decline is projected to occur.
2–4.5 mm wide, 1.75–4.5 mm high in mature cone, IUCN: EN [B1 ab(iii)+2ab(ii,iii)]
more or less convex with entire margin. Seed cones
globose, ca. 6 cm long, 5 cm wide; seed scales bossed
Uses
to shortly rostrate: mature cones not smoothly glo-
bose. Cone scales 16–20 mm long, 22–26 mm wide, Due to its usually low stature and crooked habit, this
with a rounded, thick, more or less shortly rostrate species is not exploited for timber, although an oc-
(bossed) upper margin and thin, rounded lateral casional tall specimen may have been logged from
margins ending in a cuneate base. Seeds ovoid, 9–11 forest habitat. It is not known to be in cultivation
mm long, 7–8 mm wide, with truncate base and outside a few collections in botanic gardens in the
apex; wings of unequal size; largest wing 15–20 mm (sub-)tropics.
long and 9–12 mm wide and extended at a right an-
gle to the seed; smaller wing opposite with a round-
ed edge, ca. 3 mm long and 5 mm wide.
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey, smallest wing a small acute triangular point 3–4 mm
Syn. Queensland Fl.: 498. 1883. long, opposite largest wing.

Etymology Distribution
The species epithet ‘robust’ may refer to the leaves, Papuasia: New Britain, Papua New Guinea; Austral-
which are often larger than those of the few species ia: coastal Queensland.
known in 1857 when C. Moore coined the name. TDWG codes: 43 BIS NWG-PN 50 QLD-QU

Vernacular names Ecology

165
Queensland kauri pine, Smooth-barked kauri; Agathis robusta is an emergent tree in lowland sub-
asong, muwaka, ogapa (Papua New Guinea, mostly tropical and tropical evergreen or semi-evegreen
for subsp. nesophila). rainforest. In Queensland this species (subsp. robus-
ta) occurs at its southernmost locality in a seasonal
climate with occasional winter frost, at altitudes
Description
from near sea level to 900 m a.s.l. Here the forest is
Trees to 50 m tall and 2 m or more d.b.h. with clear semi-evergreen and dominated by Eucalyptus pilu-
cylindrical bole to 30 m and a rounded or open laris (on Fraser Island). In northern Queensland it
crown with spreading to ascending branches. Bark occurs in the notophyll vine forest, a rainforest type
smooth, often lenticellate, exfoliating in roundish rich in lianas and with much higher rainfall (2000–
small scales of up to 5 cm across; outer bark brown, 3000 mm annually) than in the south. In New
reddish under newly fallen scales; inner bark pink- Guinea A. robusta is mostly represented by subsp.
ish, exuding white resin. Leaves subopposite, thin nesophila, but some specimens have been identified
or thick, multinerved, coriaceous, glabrous, light or as subsp. robusta, apparently from similar habitat.
mid lustrous green above, dull green below. Leaves Here the species is a scattered rainforest emergent
on saplings and young trees and/or in shade broadly of lower montane distribution at altitudes between
lanceolate, 8–12(–14) cm long, 3–5 cm wide, some- 600 m and 1900 m a.s.l.; in New Britain it has been
times acuminate; leaves in crowns of mature trees found at around 400 m a.s.l. It grows on a variety
narrower, lanceolate, oblanceolate or elliptic, 5–11(– of nutrient-poor soils derived from granite or other
14) cm long, 1–3 cm wide, distinctly petiolate and igneous rock, including ultrabasic types.
with an obtuse or rounded apex. Pollen cones axil-
lary, solitary on a 2–10(–18) mm long, stout pedun-
Uses
cle, cylindrical, when full grown 4–6(–7) cm long,
8–13 mm diam., becoming flexible, with 8–10 decus- The Queensland kauri pine is a valuable timber tree
sate (in 4–5 pairs), imbricate but spreading, 3–6 mm and was exploited in the past in Queensland, but
wide bract scales at their base. Microsporophylls in logging of natural stands has all but ceased there. In
tesselate (or very weakly imbricate) arrangement; New Guinea the timber of this species is used lo-
head obtuse-triangular to rhombic in outline, with cally for house construction and the resin is tapped
a raised, prismatic, nearly flat or keeled central part for various uses, but since this tree occurs very scat-
1–2 mm wide, 0.7–1.5 mm high in mature cones; tered in the forest it is not of economic importance
margins partly hidden, bearing 2–8 pollen sacs. to the timber industry. Export of round logs of Aga-
Seed cones solitary on thick peduncles, subglobose this from Papua New Guinea is banned. This spe-
or obovoid, 8–15 cm long, 8–10 cm wide, smooth, cies is widely planted as an ornamental in the tropics
green and resinous, ripening brown. Cone scales and subtropics and is present in most (sub)tropical
with slightly rounded, incurved upper margins, ca. botanic gardens and many parks. Most of the prov-
3 cm long, 3.5–4.2 cm wide, roughly triangular with enance of these trees will be populations in Queens-
rounded corners and more or less flanged on either land, especially from the northern population in
side. Seeds 12–14 × 7–8 mm, ovoid, flattened, with Cook and Kennedy Districts. It is also planted lo-
two unequal wings; largest wing 20–25 × 12–15 mm; cally with more utilitarian aims, especially resin tap-
ping in PNG.
 2 subspecies are recognized:
Conservation
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey The scattered nature of the distribution of this sub-
subsp. robusta. Dammara robusta C. Moore ex species renders commercial logging less likely; the
F. Muell., Quart. J. Trans. Pharm. Soc. Victoria 2: timber is only used locally. The main threat would
173. 1860. Type: Australia: Queensland, Cairns, be deforestation for agriculture, which may occur in
Wide Bay, C. Moore s.n. (holotype MEL?, isotype some locations with this tree, especially at lower al-
K). titudes in valleys near villages.
IUCN: NT
Description
166
Pollen cones when full grown up to 9 mm wide; Agathis silbae de Laub., Phytologia 61: 448. 1987.
micro­sporophylls obtuse-triangular, with raised Type: Vanuatu: Santo/Malo, Espiritu Santo,
head, ca. 1 × 1 mm. Tasmaloum, M. Askin 13156 (holotype NY).

Distribution Etymology
Australia: coastal Queensland (disjunct); Papua New This species was named after John Silba, an Ameri-
Guinea: Eastern Highlands, Owen Stanley Range. can with a long record of naming conifers himself.
TDWG codes: 43 NWG-PN 50 QLD-QU
Vernacular names
Conservation
No common names have been recorded for this spe-
It is not known how much effect historical log- cies.
ging has had on the size of the two populations in
Queensland, if any. Presently, logging has all but
Description
ceased and the subspecies is effectively not threat-
ened with extinction. Trees to 35(–40) m tall and 1–4 m or more d.b.h.
IUCN: LC with a clear but usually short (<10 m) bole and an
ultimately broad crown formed by long, spreading
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey and ascending branches. Bark smooth or scaly, ex-
subsp. nesophila Whitmore, Pl. Syst. Evol. 135 (1–2): foliating in large patches; colour grey weathering to
64. 1980. Type: Papua New Guinea: Morobe, Wau, white; inner bark red to pink near the wood; resin
Middle Creek, J. J. Havel s.n. (holotype K). white turning yellowish in time. Leaves subopposite,
thick, multinerved, coriaceous, glabrous, light green.
Agathis spathulata de Laub., Fl. Malesiana, ser. 1, 10 Leaves on saplings and young trees and/or in shade
(3): 435. 1988. broadly lanceolate, rarely slightly falcate, 5–12 cm
long, 2–5 cm wide, with acute or obtuse apex; leaves
in crowns of mature trees lanceolate, ovate-lanceo-
Description
late to ovate, 4–8 cm long, (1–)2–4 cm wide with an
Pollen cones when full-grown 9–13 mm wide; micro­ obtuse or acute apex. Pollen cones axillary, solitary
sporophyll heads rhombic in outline, not raised, 1.5– on a 0–6 mm long peduncle, cylindrical, when fully
2 mm wide, more or less flat or keeled. mature ca. 3.5–6 cm in length and 15–18 mm diam.;
microsporophyll arrangement markedly tessellate
when immature and becoming imbricate towards
Distribution
anthesis. Basal bract cluster loose, usually narrower
Papua New Guinea: Eastern Highlands, Owen than pollen cone, usually consisting of two or three
Stanley Range, New Britain. opposite pairs of large triangular bracts, long-bracts
TDWG codes: 43 BIS NWG-PN not present. Microsporophylls in tesselate arrange-
ment, their pedicel 2.5–2.8 mm long, slender; mi-
crosporophyll head, prismatic, 1.8–2.5 mm wide,
2.1–2.5 mm long, in adaxial view head thick with ter, western and northwestern slopes of the central
marginal flange very narrow or absent; margins en- mountain range, at altitudes between 450 m and 760
tire, slightly undulate; apex notched; head with pris- m a.s.l. No detailed record of its habitat is available,
matic process both at apex and (in micro­sporophylls but it is assumed to occupy a similar niche in the
near pollen cone apex) between base and apex. Seed forest as A. macrophylla in Fiji. Trees are various-
cones solitary on thick peduncles, globose, ca. 10–12 ly reported to grow on ridges or in hollows on the
cm diam., smooth, green, often resinous, ripening middle to upper slopes of the mountains. Very large
brown. Cone scales with rounded, incurved up- trees have been found, so it can be assumed to be
per margins, ca. 3.5 cm long, 4–5 cm wide, roughly long-lived.
triangular with rounded corners and more or less
flanged on either side. Seeds 12–15 × 7–8 mm, ovoid- 167
Conservation
oblong, with two unequal wings; largest wing 20–25
× 13–17 mm; smallest wing a small blunt triangle 4–6 The occurrence of large trees of Agathis in the interior
mm wide, opposite largest wing. mountains of the island of Espiritu Santo in Vanuatu
has long been known, both to native islanders and
at least since 1963 to visitors. De Laubenfels & Silba
Taxonomic notes
(1987) reported sightings of big emergent trees in the
This species was assumed to be identical with forest by others, but collections could not be made
A. macrophylla by Whitmore (1980), who had seen and the new species was described from a cultivated
a specimen at K from the Cumberland Peninsula tree on the coast. In April 1987 G. Bourdy collected
which has foliage, fragments of a seed cone and numerous herbarium specimens from ‘Santo Peak’
fragments of pollen cones. Later, he identified it = Mt. Tabwémasana, where the trees were observed
as A. silbae when this species had been published. to be abundant, with several very large individuals
The main distinction between the two species is seen. This was thought to be the only station of the
in the morphology of the microsporophylls of the species, but an earlier collection (specimen at K)
pollen cone, which in A. silbae are distinctive with dating from February 1979 was made at the nothern
a ‘double prismatic’ head not seen in other Agathis end of the mountain range on the Cumberland Pe-
species. The leaves, while variable, are less so than ninsula on the “top western slopes” (ca. 600 m a.s.l.)
in A. macrophylla and differ very little in shape be- and so there are at least two (sub)populations of this
tween young trees and old trees and only slightly species. There is little incentive to logging because
more in size, with much overlap. As far as is known, the large trees all have very short boles and massive
no other species is indigenous to Espiritu Santo, but branching rendering them of limited use as timber.
it may be that A. macro­phylla is present among the IUCN: VU (D2)
planted trees on the island. These planted trees are
so far restricted to village grounds near the coast.
Uses
The timber of this species is similar to that of A. mac-
Distribution
rophylla in its properties, but the shape of most trees
Vanuatu: Espiritu Santo (Cumberland Peninsula in the wild does not make them valuable as timber
and Mt. Tabwémasana = ‘Santo Peak’). trees. The inflammable white resin (‘latex’) was tra-
TDWG codes: 60 VAN ditionally used instead of kerosene for lighting fires.
Trees are grown by local people in ocean-side villag-
es of southwest Espiritu Santo, especially Tasmalum
Ecology
and Wailapa and some will have attained harvest-
Agathis silbae occurs as an emergent large tree able dimensions.
in tropical lower montane rainforest on the wet-
 Amentotaxus Pilg., Bot. Jahrb. Syst. 54: 41. 1916. Type: Amentotaxus argotaenia
(Hance) Pilg. [Podocarpus argotaenia Hance] (Taxaceae).

Latin: amentum = string; referring to the racemose white in all species, but will turn brownish in some
arrangement of pollen cones; Taxus is the classical when leaves are older than one year.
Latin name for yews.
1a. Stomatal bands as wide as the green leaf mar-
gins, or nearly so A. argotaenia
Description
1b. Stomatal bands 1.5–3(–4) times as wide as the
168 Dioecious, evergreen shrubs or small trees. Resin green leaf margins 2
canals (1) in leaves only. Bark smooth, exfoliating 2a. Stomatal bands 2–3(–4) times as wide as the
in thin flakes. Foliage shoots with conical (sub-) green leaf margins, white at first, turning
terminal buds; branching in trees rhytmic and pla- brown 3
giotropic (Massart’s model). Leaves usually opposite- 2b. Stomatal bands 1.5–2 times as wide as the green
decussate, pectinately arranged by twisted petiolate leaf margins, white or (creamy) grey, not turn-
leaf bases, flattened, large, narrowly lanceolate to ing brown 4
linear-lanceolate or slightly falcate, coriaceous, with 3a. Leaves spreading at angles of 70–90° from the
two conspicuous, broad, white stomatal bands on shoot. Pollen cones with 10–15 pairs on ca.
the abaxial side (turned downwards by twisting of 10 cm long racemes A. hatuyenensis
petioles of half of the number of leaves on a shoot; 3b. Leaves spreading at angles of 50–70° from the
in the other half the abaxial side is the lower side shoot. Pollen cones with 12–20 pairs on 10–15
without a twist). Pollen cones aggregated in umbel- cm long racemes A. yunnanensis
late clusters of (1–)3–6 racemes from large buds at or 4a. Leaf texture on the adaxial (upper) surface
just below apex of vegetative shoots, forming rows of smooth. Microsporophylls 6–8 per pollen cone,
mostly opposite pairs on each raceme, subglobose; with 2–4 pollen sacs A. assamica
microsporophylls 6–10 per cone, peltate, bearing 4b. Leaf texture on the adaxial (upper) surface ru-
2–8 small pollen sacs containing spherical pollen. gose. Microsporophylls 8–11 per pollen cone,
Seed-bearing structures usually in groups near apex with (3–)4–8 pollen sacs 5
of vegetative shoots, long pedicellate, consisting of 5a. Leaf apices long acuminate; stomatal bands
several pairs of decussate, imbricate bracts and a about twice as wide as the green leaf margins
single, terminal, erect ovule partly enclosed by the A. formosana
upper bracts and subtended by a cupular aril. Aril 5b. Leaf apices long acute or obtuse; stomatal
surrounding ripe seed greatly enlarged, ellipsoid bands about 1.5 times as wide as the green leaf
or ovoid, fleshy and succulent, red or purple. Seeds margins A. poilanei
much smaller, with a mucronate apex.

6 species. Amentotaxus argotaenia (Hance) Pilg., Bot. Jahrb.

Syst. 54: 41. 1916.
Distribution
Etymology
China, Taiwan, NE India (Arunachal Pradesh), Vi-
etnam. The species epithet (Greek argyr- = silver, and tae-
nia- = ribbon or band) describes the two stomatal
bands of the leaves.
Key to the species of Amentotaxus
The size and shape of leaves, used as characters to
Vernacular names
distinguish species by Li (1952) turn out to be vari-
able and not very reliable as diagnostic characters. Catkin yew; sui hua shan (Chinese)
The colour of the stomatal bands is more or less
 Description Ecology
Shrubs or small trees to 7 m tall. Bark smooth, ex- Amentotaxus argotaenia is widespread and occurs
foliating in thin flakes. Branches few, spreading or on limestone as well as sandstone, shale or granite,
ascending. Foliage branchlets opposite, spreading and in ravines, on steep slopes or cliffs, on summits
at 25–70° from the leading shoot axis, ascending and ridges and in mountain forests along shady
or spreading, subterete, with alternatingly twisted stream banks. The altitudinal range is between 600
grooves running from one leaf base to the next m and 1100 m a.s.l. Associated trees and shrubs
above, green in the first year, turning from green- vary with the types of rock, mainly between lime-
ish yellow to orange-brown in following years, ter- stone and other rocks. On limestone, it may grow
minating in a conical bud with ovate-triangular, with Pinus kwangtungensis, Nageia spp., Podo­carpus 169
acute scales. Leaves spreading distichously at 45–90° neriifolius, P. pilgeri, P. macrophyllus and Taxus
(–95°) from shoot axis, linear or linear-lanceolate, chinensis. On acidic rock types it is associated with
(2–)3–9(–11) cm long, falcate or nearly straight, 5–11 Amentotaxus yunnanensis, Cephalotaxus spp. and
mm wide, subsessile to short petiolate and curved broad-leaved trees and shrubs (angiosperms) like
or asymmetric at base, tapering to an obtuse or Magnolia, Quercus, and Rhododendron in montane
acute-acuminate apex; margins flat or slightly revo- evergreen or semi-deciduous forests.
lute. Leaf texture coriaceous, with sclereids causing
a mottled, rugose upper surface, dark green; on the
Uses
lower surface two white bands bordered by wide
green leaf margins and a midrib. Midrib promi- The wood is used for tool making, furniture and
nently raised on the adaxial (upper) side, lying in a wood turning (handicrafts). This species is in culti-
shallow groove, 0.8–1 mm wide, continuous to the vation in China and was introduced to Europe from
apex; on the abaxial side nearly flat and 1.2–1.5 mm Hong Kong; it makes an attractive foliage shrub but
wide. Stomata on the abaxial surface in two bands only grows outside in warmer regions. This (and
about as wide or slightly narrower than the green other) species is rare in gardens due to poor avail-
leaf margins, numerous, randomly dispersed. Pollen ability in the horticultural trade. In China it is used
cone racemes from large axillary or sub-terminal as a bonsai plant. The seeds have high oil content
buds with 2–5(–10) together, 3.5–6.5 cm long, each and are surrounded by a very striking red aril. Re-
with 10–12 cone pairs when full-grown; cones ovoid, cent investigations have been undertaken to analyse
3–3.5 mm long; microsporophylls 6–8, peltate, each its potential for anti-cancer drugs similar to those
with 2–5 pollen sacs. Seed-bearing structures near found in Taxus.
ends of foliage shoots, axillary, solitary on a slender,
down-curved, with 1.5–2 cm long peduncle, with 2 varieties are recognized:
6–8(–10) decussate, keeled bracts enclosing a single,
terminal ovule. Aril surrounding the seed ellipsoid Amentotaxus argotaenia (Hance) Pilg. var.
or narrowly obovoid, 20–26 × 10–13 mm, smooth, argotaenia. Podocarpus argotaenia Hance, J. Bot. 21:
lustrous bright red when ripe, with a mucronate 357. 1883; Nageia argotaenia (Hance) Kuntze, Revis.
apex. Seed proper much smaller, ca. 15 × 7 mm, ob- Gen. Pl. 2: 800. 1891; Cephalotaxus argotaenia
long or ellipsoid, with a small mucronate apex. (Hance) Pilg., in Engler, Pflanzenr. IV.5 [18]: 104.
1903. Type: China: Fujian, [“in jugo Lo-fau shan,
prov. Cantonensis”], E. Faber s.n. [Acc. No. 22121],
Distribution
Sep 1882, (holotype BM).
China: Chongqing, Fujian, S Gansu, Guangdong,
Guangxi, Guizhou, W Hubei, Hunan, Jiangsu, NW Amentotaxus cathayensis H. L. Li, J. Arnold Arbor.
Jiangxi, Central and SE Sichuan, SE Xizang [Tibet], 33: 195. 1952; Amentotaxus argotaenia (Hance) Pilg.
S Zheijiang; Taiwan; Indochina: Laos, N Vietnam. var. cathayensis (H. L. Li) P. C. Keng, [Chin. title; see
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- Fl. Reipupl. Pop. Sin. 7: 452. 1978]: 2. 1957.
SC CHN-GS CHS-FJ CHS-GD CHS-GX CHS-HN CHS-
JS CHS-JX CHT 38 TAI 41 lao VIE
 Description Ecology
Leaves 3–11 cm long, 6–11 mm wide. Pollen cone This variety occurs on limestone karst mountains
racemes grouped with 2–4(–5) together, rarely soli- on steep rocky slopes and ridges at around 900 m
tary. Seeds including the aril 20–26 mm long and altitude.
10–13 mm wide.
Conservation
Distribution
Amentotaxus argotaenia var. brevifolia is known
China: Chongqing, Fujian, S Gansu, Guangdong, from a very limited area and has presumably been
170 Guangxi, Guizhou, W Hubei, Hunan, Jiangsu, NW reduced in population size due to deforestation on
Jiangxi, Central and SE Sichuan, SE Xizang [Tibet], the more accessible slopes.
S Zheijiang; Taiwan; Indochina: Laos, N Vietnam. IUCN: VU (A1c)
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
SC CHN-GS CHS-FJ CHS-GD CHS-GX CHS-HN CHS-
Uses
JS CHS-JX CHT 38 TAI 41 LAO VIE
This variety grows to a small tree; its uses are similar
to those of the species but it is not known if it is in
Conservation
cultivation in or outside China.
Despite the wide distribution of the type variety of
Amentotaxus argotaenia, it is in danger of extinction
due to a similarly widespread reduction and deterio- Amentotaxus assamica D. K. Ferguson, Kew Bull.
ration of its habitat. In such cases, we have no direct 40 (1): 115. 1985. Type: India: Arunachal Pradesh,
evidence of decline but this is inferred from circum- Delei River, Camp Chibaon, F. Kingdon Ward 8026
stantial evidence. When forests on more accessible (holotype K).
mountain slopes have been and are being degraded
or even removed to make way for agriculture, this
Etymology
tree will have gone with its habitat from those lo-
calities. An assessment of deforestation will amount The species epithet refers to Assam, its native region
to an inferred assessment of the status of a forest (now Arunachal Pradesh).
­species.
IUCN: VU (A1c)
Vernacular names
Amentotaxus argotaenia (Hance) Pilg. var. brevi- Assam catkin yew
folia K. M. Lan & F. H. Zhang, Acta Phytotax. Sin.
22 (6): 492. 1984. Type: China: Guizhou, Libo,
Description
Dongjiang Shan, K. M. Lan & F. H. Zhang 82-002
(holotype GACP). Trees to 20 m tall; trunk sometimes divided low
above the ground. Bark smooth, exfoliating in thin
flakes, whitish grey. Branches spreading or sweep-
Description
ing down. Foliage branchlets opposite, spreading
Leaves 2–3.7 cm long, 5–7 mm wide. Pollen cone at 25–70° from the leading shoot axis, ascending or
racemes grouped with up to 10 together, 1.5–5.5 cm spreading, subterete or angular, with alternatingly
long. Seeds including the aril ca. 20 mm long and twisted grooves running from one leaf base to the
10 mm wide. next above, green in the first year, turning from
greenish yellow to yellowish brown in following
years, terminating in a conical bud with ovate-tri-
Distribution
angular, acute-mucronate scales. Leaves spreading
China: SE Guizhou (Libo Xian). distichously at 45–80° from shoot axis, linear or
TDWG codes: 36 CHC-GZ linear-lanceolate, (2–)7–11(–15) cm long, falcate or
more or less S-shaped, (4–)7–12 mm wide, short pet-
iolate and curved or asymmetric at base, gradually himalayana, Betula alnoides, Carpinus viminea and
tapering to an acute-acuminate apex, green with two Exbucklandia populnea. Amentotaxus assamica is a
white or silvery grey bands bordered by wide green rare species only known from two localities.
leaf margins and a midrib on the lower surface; mar-
gins flat or slightly revolute. Leaf texture coriaceous,
Conservation
with a smooth or finely striated upper surface (lack-
ing sclereids in the leaf anatomy). Midrib promi- This species is known from two disjunct localities in
nently raised on the adaxial (upper) side, lying in a Arunachal Pradesh, NE India, in only one of which
shallow groove, 0.4–0.7 mm wide; continuous to the it has been collected recently (1980s). It is obviously
apex, slightly raised and 1–1.8 mm wide on the abax- rare because several searches during the 1970s for it
ial side. Stomata on the abaxial surface in two bands remained unsuccessful until it was found again in 171
1.5–2 times as wide as the green leaf margins, nu- 1984. However, the region remains poorly explored
merous, randomly dispersed. Pollen cone racemes botanically and we cannot say whether it is restricted
from large axillary or sub-terminal buds with 2–4 to the localities known or more widespread. So far it
together, 4–5.5 cm long, each with 8–10 cone pairs is not known from outside the state. Activities like
when full-grown; cones ovoid, 3–5 mm long; micro- road building are opening up this border region to
sporophylls 6–8, peltate, each with 2–4 pollen sacs. settlement, leading to shifting agriculture and con-
Seed-bearing structures axillary, solitary on 1.5–2.5 sequently forest disturbance and destruction. There
cm long peduncles, with 8 decussate, ovate, keeled, is an urgent need to establish forest reserves as well
2.5–5 mm long and 2.5–5 mm wide bract scales (the as cultivation action to provide ex situ conservation
upper scales largest). Aril obovoid-oblong to ellip- for this species (Sahni, 1990).
soid, 25–35 × 15–25 mm, lustrous green turning yel- IUCN: VU (A1c)
low to purple, with a mucronate apex. Seed proper
much smaller, 15–20 × 7–10 mm, oblong or ellipsoid,
Uses
with a small mucronate apex.
No specific uses have been recorded of this species.
Taxonomic notes
Ferguson (1985) has distinguished this species from Amentotaxus formosana H. L. Li, J. Arnold Arbor.
A. yunnanensis mainly on leaf anatomical characters. 33: 196. 1952. Amentotaxus yunnanensis H. L. Li var.
One of these, the lack of sclereids in the upper meso- formosana (H. L. Li) Silba, Phytologia 68: 25. 1990.
phyll of the leaves, present in all other species, gives Type: Taiwan: Pingtung Co., [“Taririku”], S. Sasaki
the leaves a smooth texture quite different from its s.n. (holotype TAI). Fig. 38, 39
congeners. At the time, the seed bearing structures
were unknown, these were later described from
Etymology
newly collected specimens and appear to be larger
than in most other species, though variable. The species epithet refers to Taiwan (formerly For-
mosa), where this species is indigenous; see also
Latin: formosus = handsome or well formed.
Distribution
India: Arunachal Pradesh [“Assam”].
Vernacular names
TDWG codes: 40 EHM-AP
tai wan sui hua shan (Chinese)
Ecology
Description
Amentotaxus assamica occurs in warm temperate
montane rain forest on N-facing slopes at altitudes Small trees to 10 m tall. Bark smooth, brown, ex-
between 1600 m and 2000 m a.s.l. These are mossy foliating in thin flakes. Branches few, spreading or
forest dominated by Quercus spp., Castanopsis spp., ascending. Foliage branchlets opposite, spreading
Acer sp. and Rhododendron spp., with associated at 25–70° from the leading shoot axis, ascending
taxa such as Magnolia sp., Michelia sp., Corylopsis or spreading, subterete, with alternatingly twisted
 grooves running from one leaf base to the next
Distribution
above, green in the first year, turning to yellowish
brown with dark grooves in following years, termi- Taiwan (southernmost mountains).
nating in a conical bud with ovate-triangular, acute TDWG codes: 38 TAI
scales. Leaves spreading distichously at 50–70° from
shoot axis, lanceolate or linear-lanceolate, (3.5–)5–
Ecology
8.5(–9.5) cm long, slightly or more strongly falcate,
5–10 mm wide, subsessile to short petiolate and Amentotaxus formosana is a rare tree in montane
curved or asymmetric at base, gradually tapering to evergreen tropical rainforest and broad-leaved sub-
a long acuminate apex, dark green, with two white tropical forest. It is usually found on steep slopes,
172 bands bordered by wide green leaf margins and a in ravines, or on cliffs in the sub-canopy under
midrib on the lower surface; margins (slightly) revo- taller trees. Its altitudinal range is between 500 m
lute. Leaf texture coriaceous, with sclereids causing and 1300 m a.s.l. with most trees occurring above
a mottled, rugose upper surface. Midrib promi- 900 m a.s.l. In these primary forests members of
nently raised on the adaxial (upper) side, lying in a the Fagaceae (Castanopsis, Lithocarpus, Quercus)
shallow groove, 0.8 mm wide, continuous to apex, are often the dominant or emergent trees and there
raised at least near base and 1–1.5 mm wide on the is an abundance of ferns, including tree ferns, and
abaxial side. Stomata on the abaxial surface in two shrubs. Podo­carpus nakaii is another conifer that
bands about twice as wide as the green leaf margins, occurs here.
numerous, randomly dispersed, thickly covered
with white wax. Pollen cone racemes from large ax-
Conservation
illary or sub-terminal buds with 2–5 together, rarely
solitary, 1.5–3.5 cm long, each with 7–10 crowded This species was evaluated as Endangered (EN) by
cone pairs when full-grown, cones subglobose, ca. the Taiwanese botanist S. Y. Lu in Rare and Endan-
2.5 mm long; microsporophylls 8–10, peltate, each gered Plants in Taiwan 1 (1996) and as Critically
with 5–8 tiny pollen sacs. Seed-bearing structures ­Endangered (CR) by the Conifer Specialist Group
near ends of foliage shoots, axillary, solitary on a in the Status Survey and Conservation ­Action
slender, down-curved, 1.5–2 cm long peduncle, with Plan: Conifers (Farjon & Page, 1999), both under
8–10 decussate, keeled bracts enclosing a single, ter- the now superseded 1994 criteria. This species is
minal ovule. Aril surrounding the seed ellipsoid or certainly rare, but is also very difficult to find (as I
narrowly obovoid, 20–25 × 9–11 mm, smooth, lus- experienced recently) and deciding on the number
trous reddish purple when ripe, with a mucronate of mature individual plants under the (amended) C-
apex. Seed proper much smaller, ca. 15 × 7 mm, ob- criterion (250 for CR, 2500 for EN) is much more
long or ellipsoid, with a small mucronate apex. difficult than it sounds. Assuming that more than
250 trees still exist, Lu’s estimate is probably more
correct, also because it has recently been found in
Taxonomic notes
‘new’ localities. The greatest threat to it appears to be
This species was formerly not recognized as being conversion of native mixed evergreen forest to plan-
distinct from A. argotaenia and many earlier ac- tations with primarily Cryptomeria japonica, which
counts of the flora of Taiwan refer to it under this has been pursued at least until recently, when the
name. In fact, A. argotaenia appears to occur in government vowed to discontinue this programme
Taiwan as well, making things not easier. The main of ‘afforestation’ in favour of natural regeneration
distinction between the two species is in the width where possible. This species is present in the Tawu
of stomatal bands, which in A. formosana are about Taiwan Nature Reserve, a small forest reserve of
twice as wide as the green leaf margins. With leaves 86 ha.
about equally wide, this amounts to narrower leaf IUCN: EN C2a (i)
margins in A. formosana compared with leaves of
A. argotaenia. The pollen cone racemes of A. for-
Uses
mosana are shorter than those of A. argotaenia,
resulting in crowded pollen cones in A. formosana The wood, though of small dimensions, is valued and
where they are spaced in A. argotaenia. used for the making of furniture, farm implements,
tools and utensils, and wood turning for souvenirs. each with 10–15 cone pairs when full-grown, cones
This species is in cultivation as an ornamental tree subglobose, ca. 3.5 mm long; microsporophylls 8–10,
in China, Japan and Taiwan, but very rare and only peltate, each with 5–8 tiny pollen sacs. Seed-bearing
seen in some botanic gardens elsewhere. Its broad, structures near ends of foliage shoots, axillary, soli-
white bands of stomata are very striking, although tary on a 1.5 cm long peduncle, with 10–12 decussate,
normally hidden from view on the underside of keeled bracts enclosing a single, terminal ovule. Aril
leaves. Lack of commercially available cuttings or surrounding the seed only known in immature state.
seed has prevented this species from being grown
more widely; it is obviously not suitable for climates
Taxonomic notes
with winter frost.
This species remains taxonomically poorly known; 173
its main distinguishing character from Amentotaxus
Amentotaxus hatuyenensis Hiep, Fl. Cambodge, yunnanensis seems to be the colour of the stomatal
Laos et Vietnam 28: 126. 1996. Type: Vietnam: bands on the abaxial leaf surface. These bands are
Ha Tuyên Prov., Pho Bang, Lung Cun, Vu Xuan described in the original description as “rousses ou
Phuong 329 (holotype P). brunâtres”, but in A. yunnanensis they are also not
entirely white but become yellowish white or pale
brown. Initially, in new leaves at least, all stomatal
Etymology
bands seem to be white, they may turn to brown
The species epithet refers to Ha Tuyên Province, later or only in drying leaves. The leaf shape, at least
from where it was first described. in the type specimen of A. hatuyenensis, is more
lanceolate than in A. yunnanensis, but there was no
other material available when A. hatuyenensis was
Vernacular names
described and we still know little about variation in
De tung soc nau rong (Vietnamese) both characters. I have not seen the later specimens
from Ha Giang Province ascribed to this species
and cannot say how much they differ from A. yun-
Description
nanensis. More recently, trees or shrubs found in
Shrubs to 4–5 m tall. Foliage branchlets slender, some localities in Ha Giang Province reported to be
terete, with grooves alternately running from one A. hatuyenensis have turned out to be either A. yun-
leaf base to the next, green turning yellowish or pale nanensis or A. argotaenia (Philip Thomas, RBGE,
brown, terminating in a conical bud with ovate-­ pers. comm. May 2008). The characters mentioned
triangular, acute scales. Leaves spreading distichous- may only indicate two varieties, not two species.
ly at 70–90° from shoot axis, lanceolate, 4–7 cm long,
usually straight or sometimes slightly falcate, 7–13
Distribution
mm wide, subsessile to short petiolate and broadly
cuneate or obtuse at base, tapering to an acuminate Vietnam (Ha Giang Prov., Ha Tuyên Prov.).
apex, light green or mid green, with two initially TDWG codes: 41 VIE
white or yellowish white, later reddish brown or
brown bands bordered by wide green leaf margins
Ecology
and a midrib on the lower surface; margins revolute.
Leaf texture coriaceous, with sclereids causing a This is a rare species occurring on steep ridges of
mottled, rugose upper surface. Midrib prominently limestone karst mountains at altitudes between 1000
raised on the adaxial (upper) side, lying in a shal- m and 1500 m a.s.l. In its few known localities it may
low groove, 0.8–1 mm wide and continuous to the grow together with the conifers Pinus fenzeliana,
apex, raised and 1–1.5 mm wide on the abaxial side. Tsuga chinensis, Amentotaxus yunnanensis, Cepha-
Stomata on the abaxial surface in two bands 2–3(–4) lotaxus mannii, Xanthocyparis vietnamensis, Nageia
times as wide as the green leaf margins, numerous, fleuryi, Podocarpus neriifolius and P. pilgeri, as well
randomly dispersed, white or cream in a brown as with various angiosperm trees, e.g. Acer, Carpi-
matrix. Pollen cone racemes from large axillary or nus, Lithocarpus, Quercus, and Ulmus and a rich epi-
sub-terminal buds with 3–5 together, ca. 10 cm long, phytic flora of mosses, ferns, and orchids. Frequent
 fog and rain amount to annual precipitation above in flakes. Branches spreading or ascending. Foliage
1800 mm, usually under cool temperatures of ca. branchlets opposite, spreading at 25–60° from the
15–18 °C. leading shoot axis, ascending or spreading, angular
in cross-section, with alternatingly twisted grooves
running from one leaf base to the next above, green
Conservation
in the first year, turning to yellowish brown in fol-
When described in 1996, this species was known lowing years, terminating in an ovoid bud with
from a single locality in Ha Tuyen Province; sub- ovate-triangular, keeled and acute scales. Leaves
sequently botanical surveys have found it in a few spreading distichously at 50–70° from shoot axis, el-
other localities, all in Ha Giang Province, one is in liptical, oblanceolate or linear, (3–)5–8(–9) cm long,
174 the Bat Dai Son Nature Reserve, known to protect straight to slightly or more strongly falcate, (4.3–)5–
the new genus and species in the Cupressaceae: Xan- 8.5 mm wide, short petiolate and more or less gradu-
thocyparis vietnamensis. Amentotaxus hatuyenen- ally narrowing at base, gradually tapering to a long
sis remains poorly known, and although currently acute or obtuse apex, dark green, with two white or
thought to be a local endemic, it may turn up in oth- creamy grey bands bordered by green leaf margins
er locations with continued surveys in future. Forest and a midrib on the lower surface; margins revolute.
degradation as a result of fires, fuelwood and tim- Leaf texture coriaceous, with sclereids causing a
ber cutting are adversely influencing its habitat, but mottled, rugose upper surface. Midrib prominently
the shrubby habit of this species makes it unlikely raised on the adaxial (upper) side, lying in a shallow
to be targeted for cutting. Observed regeneration is groove, 0.5 mm wide, continuous to apex, raised at
poor, but this is also an imperfectly known factor at least near base and 1–1.8 mm wide on the abaxial
present. side. Stomata on the abaxial surface in two bands
IUCN: EN (A2c, B1ab [iii]) about 1.5 times as wide as the green leaf margins,
numerous, randomly dispersed, thickly covered
with white wax. Pollen cone racemes from large ax-
Uses
illary or sub-terminal buds with 3–4 together, rarely
No uses have been recorded of this species. solitary, 2.5–4.5 cm long, each with 8–12 cone pairs
when full-grown, cones subglobose or ovoid, 3–4
mm long; microsporophylls 8–11, peltate, each with
Amentotaxus poilanei (Ferré & Rouane) D. K. Fer­ (3–)4–6(–8) pollen sacs. Seed-bearing structures
guson, Adansonia, sér. 4, 11 (3): 316. 1989. Amento­ near ends of foliage shoots, axillary, solitary on a
taxus yunnanensis H. L. Li var. poilanei Ferré & slender, down-curved, 1–2 cm long peduncle, with
Rouane, Trav. Lab. Forest. Toulouse T. 1 (9, 1): 3. ca. 8 decussate, keeled bracts enclosing a single, ter-
1978. Type: Vietnam: Kon Tum Prov., Ngoc Pan minal ovule; only known in immature stage.
Massif, Mt. Ngoc Linh, E. Poilane 32686 (holotype
P).
Taxonomic notes
This species was originally described as a variety
Etymology
of A. yunnanensis, based on observed variations of
This species is named after Eugène Poilane (1888– leaf length and width. As pointed out by Ferguson
1964), who collected the type specimens. (1989), who raised this taxon to species rank, leaf
dimensions are of dubious value to distinguish be-
tween species as there is usually much overlap and
Vernacular names
the measurements vary considerably within species.
Poilane’s catkin yew; De tung Nam, Sam bong Nam Ferguson found that length/width ratios are prob-
(Vietnamese) ably more informative, showing A. poilanei to have
narrower leaves. These values similarly overlap, but
to a lesser extent. He then went on to describe fea-
Description
tures of leaf anatomy of both taxa and found several
Trees to 20 m tall; trunk to 1 m d.b.h. Bark smooth, differences to justify not only to retain var. poilanei
becoming scaly on large trunks, brown, exfoliating as distinct but to raise it to species level. There is a
single resin duct in the leaf that is much narrower
Uses
in A. poilanei than in A. yunnanensis, and ‘flanges’
are absent in the revolute leaf margins of A. poilanei. No uses have been recorded of this species. The size
The morphology and anatomy of A. poilanei are, ac- of the largest trees would certainly make it valuable
cording to Ferguson, closer to A. formosana than to as a timber tree used for the making of furniture,
A. yunnanensis. Like the other localized Vietnam- tools, etc. Inaccessibility is probably a factor limiting
ese species, A. hatuyenensis, A. poilanei remains a its use. It is not known to be in cultivation at present.
somewhat doubtfully distinct species. The as yet
unknown mature female organs do not help solving
this problem, so more good material needs to be col- Amentotaxus yunnanensis H. L. Li, J. Arnold
lected from the single locality where it is known to Arbor. 33: 197. 1952. Amentotaxus argotaenia 175
occur. (Hance) Pilg. var. yunnanensis (H. L. Li) P. C. Keng,
[Chin. title; see Fl. Reipupl. Pop. Sin. 7: 452.
1978]: 2. 1957. Type: China: SE Yunnan, Maguan,
Distribution
[“Makwan”], H. T. Tsai 51887 (holotype US). Fig. 40
Vietnam (Kon tum Prov., Mt. Ngoc Linh).
TDWG codes: 41 VIE
Etymology
The species epithet refers to Yunnan, China, from
Ecology
where it was first described.
This species is apparently a large tree occurring in
high montane closed evergreen rainforest, at an al-
Vernacular names
titude around 2300 m a.s.l. It is locally common but
scattered, mixed with broad-leaved (angiosperm) Yunnan catkin yew; Yunnan sui hua shan (Chinese);
trees and perhaps Nageia wallichiana as the only De tung Van Nam (Vietnamese)
other conifer present. Rainfall is very high, at least
over 3000 mm per annum and cool temperatures
Description
prevail due to almost continuous cloud cover.
Trees to 25(–30) m tall; trunk to 80 cm d.b.h. Bark
smooth, brown, exfoliating in thin flakes. Branches
Conservation
spreading or ascending, forming a broad crown. Fo-
This species is still only known with certainty from liage branchlets opposite, spreading at 45–80° from
a single mountain, where it was discovered in 1946. the leading shoot axis, ascending or spreading, sub-
Reports from other localities need confirmation by terete, with alternatingly twisted grooves running
taxonomists who know the genus Amentotaxus well. from one leaf base to the next above, green in the
The primary forest in this locality is still present and first year, turning to light yellow or yellowish brown
the total population probably consists of fewer than in following years, terminating in a conical bud with
1000 mature trees. The status of A. poilanei was as- ovate-triangular, acute scales. Leaves spreading dis-
sessed under the IUCN Red List criteria of 1994 as tichously at 50–70° from shoot axis, linear or linear-
Vulnerable, assuming a modest decline due to for- lanceolate, (3.5–)5–10(–15) cm long, usually straight
est fragmentation at lower altitudes, approaching or or sometimes slightly falcate, 8–12(–15) mm wide,
encroaching on the population. More recent visits subsessile to short petiolate and broadly cuneate
have indicated that there are no direct threats at or obtuse at base, tapering to an obtuse or more
present and that the species should be classified as or less acuminate apex, light green or mid green,
Vulnerable on the basis of its small population size with two yellowish white to pale brown bands bor-
alone, using the revised 2001 criteria. There are pro- dered by wide green leaf margins and a midrib on
tected forest areas on the mountain which include the lower surface; margins (slightly) revolute. Leaf
this species. texture coriaceous, with sclereids causing a mottled,
IUCN: VU (A2c) rugose upper surface. Midrib prominently raised on
 the adaxial (upper) side, lying in a shallow groove, mannii, Dacrycarpus imbricatus and Nageia walli-
0.8–1 mm wide, continuous to apex, raised at least chiana. It is a relatively shade tolerant species, with
near base and 1–1.5 mm wide on the abaxial side. seedlings and saplings successfully growing up un-
Stomata on the abaxial surface in two bands 2–3 der a forest canopy.
times as wide as the green leaf margins, numerous,
randomly dispersed, white or cream in a pale brown
Conservation
matrix. Pollen cone racemes from large axillary or
sub-terminal buds with 4–6 together, 10–15 cm long, Due to recent discoveries made of this species dur-
each with 12–20 cone pairs when full-grown, cones ing intensive botanical surveys in Vietnam, this spe-
subglobose, ca. 3.5 mm long; microsporophylls 8–10, cies is now known to have a wider distribution than
176 peltate, each with 4–6(–8) tiny pollen sacs. Seed- previously assumed. The majority of its (sub)popu-
bearing structures near ends of foliage shoots, ax- lations appear to be in Vietnam, not in China, and
illary, solitary on a slender, down-curved, 1.5–2 cm it is also in Vietnam where the species attains large
long peduncle, with 8–10 decussate, keeled bracts tree size. A re-assessment for Vietnam (Nguyen
enclosing a single, terminal ovule. Aril surrounding Tien Hiep et al., 2004) gave it the status of Vulner-
the seed ellipsoid or narrowly ovoid, 22–26(–30) × able (VU) for that country; the implication of this is
12–15 mm, smooth, lustrous bright red to reddish that its global status cannot be more threatened and
purple when ripe, with a mucronate apex. Seed should also become VU on re-assessment. In China
proper much smaller, ca. 18 × 10 mm, oblong or el- it has been given the status of Endangered (EN); this
lipsoid, with a small mucronate apex. has lead to a similar global status based on the ear-
lier assumption that most populations were within
that country. Its current status on the IUCN Red
Distribution
List therefore needs amendment. It is still threat-
China: Guizhou (Xingyi), Yunnan; N Vietnam (Bac ened mainly by deforestation, leading to fragment-
Can, Ha Giang, Ha Tuyen, Lao Cai, Nghe An and ed areas of forest with declining population size in
Thanh Hoa provinces); Laos (Houaphan Prov.). most. Targeted logging removes seed-producing
TDWG codes: 36 CHC-GZ CHC-YN 41 LAO VIE trees, which may lead to diminished recruitment of
seedlings and saplings in the population. Few (sub)
populations occur in protected areas and it has been
Ecology
observed that this is often insufficient to protect the
Amentotaxus yunnanensis is usually a small under- species from deforestation or logging.
storey tree occurring scattered in mixed evergreen IUCN: EN (A1c)
and deciduous broad-leaved forests, but occasional-
ly it can become much larger and reach into the can-
Uses
opy of primary closed evergreen tropical rainforest.
The altitudinal range is between 800 m and 1600 m The timber of larger trees is valued for furniture
a.s.l. Annual precipitation is above 1500 mm and fog making, while smaller sizes go to handicrafts and
is frequent. It is most commonly found on limestone tool making. Much of this is done in China and it
karst formations, where it mixes with other conifers, is likely that timber of Vietnamese origin is used.
e.g. Fokienia hodginsii, Xanthocyparis vietnamen- The seeds are rich in oils and these are extracted
sis (locally), Pseudotsuga sinensis, Tsuga chinensis, for medi­cinal purposes. Screening for anti-cancer
Podo­carpus neriifolius, Dacrydium elatum and ­Taxus drugs, as in other species like A. argotaenia, may be
chinensis, as well as with angiosperms. On substates conducted with this species as well, now that it is
derived from silicious rocks (granite, gneiss) it is known to occur more widely. This species is in lim-
sometimes a large tree amongst angiosperms, with ited cultivation, e.g. as a bonsai plant.
only a few associated conifers e.g. Cephalotaxus
 figure 2. Abies
amabilis in
Wenatchee N.F.,
Washington, USA

figure 1. Abies
alba in the Forêt de
la Joux, Jura Mts.,
France

figure 4. Abies
­cephalonica flushing
leaves

figure 3. Abies bracteata in the

Santa Lucia Mts., California, USA

figure 6. Abies delavayi var.

­delavayi seed cone

figure 5. Abies
delavayi var. delavayi
seed cones
figure 7. Abies
­delavayi var.
­nukiangensis seed
cones
 figure 8. Abies
homolepis pollen cones

figure 10. Abies

koreana young seed
cones

f i g u re 9 . Abies kawakamii stand in Taroko N.P., Taiwan

f i g u re 13 . Abies numidica tree

with seed cones

figure 11. Abies magnifica in the Sierra ­Nevada,

California, USA

figure 12. Abies nebrodensis seed cone

 figure 1 4. Abies pinsapo helically
­arranged leaves

f i gu re 15 . Abies pinsapo fig ure 16. Abies procera seed cones

seed cone (photo W. Milliken)

figure 17. Abies recurvata var. ernestii

seed cones (photo X. C. Zhang)

fi g u re 1 8 . Abies spectabilis f i g ure 1 9. Abies squamata bark of young

mature seed cone tree (photo P. de Spoelberch)
figure 2 0. Abies squamata bark of old
tree (photo P. de Spoelberch)
figure 22. Actinostrobus arenarius along Hwy. 123, Western Australia

f i gu re 2 1 . Acmopyle pancheri foliage and seed

cones (© Bedgebury Pinetum)
 f i g u re 2 4. Afrocarpus falcatus foliage and seeds
(photo D. Luscombe)

fi g u re 23 . Actino­strobus f i g u re 2 6 . Afrocarpus gracilior foliage

arenarius seed cones and seeds (photo Forest & Kim Starr)

f i g ure 27. Agathis australis in Trounson

Park, North Island, New Zealand

figure 2 5. Afro-
carpus gracilior tree
in Ethiopia (photo
J. Grimshaw)

figure 2 8. Agathis
australis The Tane
Mahuta (Lord of
the Forest)

figure 30. Agathis

borneensis leaves

figure 29. Agathis

australis seed cones
 figure 32. Agathis kinabaluensis sapling
on the Mesilau River

figure 31. Agathis

­kinabaluensis on the Mesilau
River, Mt. Kinabalu, Borneo

figure 37. Agathis ovata

leaves and pollen cone

f i g u re 3 3 . Agathis lenticula tree at figure 35. Agathis microstachya trees at Lake Barrine, Queensland
­Kinabalu Park H.Q., Borneo
figure 36. Agathis ovata trees near Yaté, New Caledonia
f i g u re 3 4 . Aga-
this lenticula trunk
in Crocker Range,
Borneo
 fig ure 3 8. Amentotaxus
formosana tree in southern
Taiwan (photo C. N. Page)

fig ure 40. Amentotaxus

yunnanensis in Ha Giang,
Vietnam (photo P. Cribb)

fi g u re 4 2 . Araucaria araucana pollen cones

fig ure 4 1. Araucaria araucana in Chile

(photo M. Gardner)

figure 39. Amentotaxus formosana

leaves underside

fig ure 43. Araucaria araucana

seed cones
fig ure 4 4. Araucaria bernieri
in New Caledonia

fig ure 45. Araucaria bidwillii emergent

trees in the Bunya Mts. Queensland
 figure 4 7. Araucaria
columnaris on the Isle of
Pines, New Caledonia

fig ure 46. Araucaria

bidwillii tree in the Bunya
Mts. Queensland

figure 50. Araucaria

heterophylla seed cones

fig ure 4 8. Araucaria

columnaris seedlings

figure 5 1. Araucaria
laubenfelsii foliage with
pollen cones

fig ure 4 9.  ­Araucaria

cunninghamii var. cunning­
hamii tree in the ­Bunya Mts.
Queensland

figure 52. Araucaria

muelleri in New Caledonia
 figure 54. Araucaria
­scopulorum foliage
fi g u re 5 3 . Araucaria scopulorum seed
cones in New Caledonia

fig ure 56. Athrotaxis selaginoides at Dove Lake, Tasmania, Australia

f i gu re 5 5 . Athrotaxis cupressoides at
Dove Lake, Tasmania, Australia

f i gu re 5 8 . Austrocedrus chilensis forest on lava, Chile (photo M. Gardner)

figure 5 7. Athrotaxis selaginoides seed cones
figure 59. Austrocedrus chilensis seed cones
Araucaria Juss., Gen. Pl.: 413. 1789. Dombeya Lam., Encycl. 2: 301. 1786, non Cav.
(1785); Columbea Salisb., Trans. Linn. Soc. London 8: 317. 1807; Eutassa Salisb.,
Trans. Linn. Soc. London 8: 316. 1807; Eutacta Link, Linnaea 15: 543. 1842. Type:
Araucaria araucana (Molina) K. Koch [Pinus araucana Molina] (Araucariaceae).

Marywildea A. V. Bobrov & Melikyan, Komaro- imbricate, spreading at 45–90° from a thin to stout
via 4: 57. 2006. Type: Marywildea bidwillii (Hook.) rachis, apically free at anthesis, divided into a stalk
A. V. Bobrov & Melikyan [Araucaria bidwillii and a lamina; stalk thin and weak, linear; lamina tri-
Hook.]. Titanodendron A. V. Bobrov & Melikyan, angular, rhombic or peltate. Pollen sacs 6–20. Seed 185
Komarovia 4: 60. 2006 Type: Titanodendron hun- cones confined to the mid and upper crown, axillary
steinii (K. Schum.) A. V. Bobrov & Melikyan [Arau- or terminal, sessile or borne on a short, robust pe-
caria hunsteinii K. Schum.]. duncle-like shoot, subtended by modified leaves and
then a transition zone of sterile cone bracts, large,
Name derived from the Araucanos (more correctly globose and ± heavy (to 5, occasionally 10 kg), with
known as Pehuenche) who live in the region of the milky resinous exudate, usually ripening in second
type species. year. Bract-scale complexes numerous, spirally ar-
ranged on a ± thick rachis. Bracts broad, obtrullate,
ovoid-oblong, cuneate or flabellate, often extended
Description
laterally into membranous (occasionally woody)
Small to very tall monoecious or more rarely dio- wings, apical margin thickened and with a narrow
ecious trees with monopodial branching; branched apophysis at tip. Seed scale enclosed within and
to ground when young but with clear bole when partly fused with bract, not or slightly wider than the
old, resinous. Trunk terete, clothed with leaves seed, terminating in a free scale-like ‘ligule’ reaching
when young. Crown columnar to candelabra- or more-or-less to base of the spur of the fertile bract.
­umbrella-shaped. Bark exfoliating in plates and Ovules one (occasionally 2) per scale. Seeds cuneate;
scales, or in horizontal strips. Branching conform- seed coat fused partly or wholly with scale, in one
ing to either Massart’s or Rauh’s model depending species (A. bidwillii) scale and seed becoming sepa-
on species (sometimes intermediate between the rated; wing absent. Germination epigeal or hypoge-
two), often reiterating. Apex of shoot a cluster of in- al; cotyledons in 2 free and 2 fused pairs, with 4 free,
completely formed leaves; bud scales absent. Leaves or 4 fused into 2 pairs at base.
persisting for many years (in some species including
on trunk). All leaves similar or more often differen- 19 species in 4 sections.
tiated into more-or-less distinct juvenile and adult
states. Leaves spirally arranged on all axes including
Distribution
the trunk, multi-veined, weakly to strongly keeled
abaxially and sometimes adaxially; apex incurved or South America: Argentina, Brazil, Chile, Para-
not; margin entire or papillate or denticulate. Stoma- guay; Australia: New South Wales, Norfolk Island,
ta amphistomatic (often very unequally, with fewer Queensland; New Caledonia; New Guinea.
stomata abaxially). Resin canals in leaves alternating
with vascular bundles and in the same plane, or pe-
Synopsis
ripheral. Pollen cones solitary, terminal or axillary
(lateral) on the ultimate foliage branchlets, on dif- Recent phylogenetic analysis (Hollingsworth et al.,
ferent branchlets to the seed cones and often lower unpubl. data RBG Edinburgh) using various DNA
down on the tree and/or on younger trees, subtend- sequence data supports a taxonomic division of the
ed by a cluster of ± modified leaves, initially erect, genus into 4 sections. These sections can also be rec-
frequently pendulous when shedding pollen, cylin- ognized by specific combinations of morphological
drical to ovoid, varying considerably in size between characters; some of which are given below with the
species and sometimes massive (up to 25 cm long), section names.
deciduous after shedding pollen. Microsporophylls
 Sect. Araucaria. Dioecious or less commonly mo- well as intermediate shapes. Seedlings have different
noecious. Leaves undifferentiated into juvenile and leaves from saplings or young trees as well as ma-
adult states, large, flat, thick or thin. Pollen cones ture trees. On mature trees adventitious branching
axillary, often 2-several close together at tips of foli- often occurs and the leaves on these branches are
age branches. Bract-scales nut-like, without wings, quite similar, but not equal to those on young trees.
indehiscent; seed retained on scale when shed. Ger- The adult leaves described in the keys are those of
mination hypogeal. Cotyledons 2, long-stalked in the ultimate primary foliage branches of mature
germination, stalks not fused. Seedling fleshy. Spe- trees that can produce cones and exclude those of
cies: A. angustifolia, A. araucana. adventitious branches. Under cultivation, trees may
retain (semi–)juvenile leaf shapes and sizes longer
186 Sect. Bunya Wilde & Eames. Monoecious. Leaves than in their natural habitat, especially when grown
undifferentiated into juvenile and adult states, large, under glass. It is virtually impossible to key these
flat, spreading or slightly imbricate. Pollen cones ax- out, as juvenile leaves of many species are highly
illary, often with 2–6 together. Seed cones axillary, similar. Pollen cones, as in the genus Agathis, have
subsessile or shortly pedunculate. Seeds only partly more informative characters than seed cones and
fused with scale, becoming separated. Germination are sometimes crucial for determination of species
hypogeal. Cotyledons 2, remaining inside the seed in addition (or in preference) to foliage. They can of-
coat. Seedling fleshy. Species: A. bidwillii. ten been found under the tree (most species are mo-
noecious) and are then full size, having shed their
Sect. Eutacta Endl. Monoecious. Leaves clearly dif- pollen, and can be used for comparison of character
ferentiated into juvenile and adult states; juvenile states if not too far disintegrated. Trees grow a trunk
leaves more-or-less narrow, falcate, spreading; adult and branches in specific ways that may be distinct
leaves scale-like and imbricate, adpressed. Pollen among taxa. The architecture of branching has been
cones terminal. Seed cones terminal, long-pedun- studied and models are named after their describers.
culate. Bract-scales samara-like, thinly winged, Two of these models apply to the species of Arau-
indehiscent; seed retained on scale when shed. Ger- caria, Massart’s and Rauh’s. In both, there is a sin-
mination epigeal. Cotyledons 4, completely free gle, dominant, erect leader making the stem (trunk
or fused into two pairs at base, freed from seed at in mature trees) and branching occurs in pseudo-
germination. Seedling not fleshy. Species: A. berni- whorls at regular intervals on it. The branches spread
eri, A. biramulata, A. columnaris, A. cunninghamii, out more or less horizontally. In Massart’s model
A. heterophylla, A. humboldtensis, A. laubenfelsii, they remain horizontal, as do the lateral branches,
A. luxurians, A. montana, A. muelleri, A. nemorosa, or they may be more or less drooping, and buds fac-
A. rulei, A. schmidii, A. scopulorum, A. subulata. ing up and down on each subsequent pseudo-whorl
remain dormant. In Rauh’s model, the branches are
Sect. Intermedia C. T. White. Monoecious. Leaves curved upward towards their ends and all buds on
differentiated into juvenile and adult states, juve- a pseudo-whorl can grow to form new branches,
nile leaves small, needle-like, flat; adult leaves large, repeating the pattern on the stem. Adventitious
flat, spreading, sometimes slightly imbricate. Pol- branching, often triggered by some damage, usually
len cones axillary. Seed cones axillary. Bract-scales does not conform to the model and can disrupt the
­samara-like, with broad relatively thin wing, in- architecture, so only primary branching should be
dehiscent; seed retained on scale when shed. Ger- considered.
mination epigeal. Cotyledons 2, free, freed from
seed at germination. Seedling not fleshy. Species: 1a. Leaves differentiated into juvenile and adult
A. ­hunsteinii. stages. Bract scales of seed cones samara-like.
Seedlings not fleshy 2
1b. Leaves undifferentiated into juvenile and adult
Key to the sections of Araucaria
stages. Bract scales of seed cones nut-like or
The leaves of many species in the genus Araucaria large and heavy. Seedlings fleshy 3
are highly variable and part of this variation is as- 2a. Adult leaves imbricate, often adpressed. Coty-
sociated with the age of the branch on which they ledons of seedlings 4, free or sometimes fused
appear. There are often juvenile and adult forms, as into two pairs at base Sect. Eutacta
2b. Adult leaves spreading. Cotyledons of seedlings × 7–9 cm; bract scales 22–25 × 15–20 mm
2, free Sect. Intermedia; a single species, A. laubenfelsii
A. hunsteinii 4b. Adult leaves broadly ovate. Crown domed, very
3a. Bract scales of seed cones nut-like, without open. Mature seed cones 8–11 × 7–9 cm; bract
wings, indehiscent; seed retained on scale when scales 25–35 × 18–25 mm A. montana
shed Sect. Araucaria 5a. Branching architecture according to Rauh’s
3b. Bract scales of seed cones large, heavy, with model, without or with infrequent adventitious
woody wings, dehiscent; seed shed from scale branching, developing an open crown in ma-
at maturity ture trees A. biramulata
Sect. Bunya; a single species, A. bidwillii 5b. Branching architecture according to Mas-
sart’s model, often with frequent adventitious 187
branching, developing a more or less dense
Key to the species of Section Araucaria
crown in mature trees 6
1a. Foliage branches with and including adult 6a. Adult leaves ovate or broadly ovate with a
leaves up to 40 mm wide. Pollen cones 15–30 rounded apex, 4–16 × 2.5–6.5(–8) mm. Pollen
mm wide. Resin exuding from bark on trunk cones (10–)12–15(–17) cm long and 20–28 mm
reddish A. angustifolia wide A. luxurians
1b. Foliage branches with and including adult 6b. Adult leaves from subulate-acicular to broadly
leaves 60–90 mm wide. Pollen cones 40–50 ovate with an acute, acuminate or obtuse, never
mm wide. Resin exuding from bark on trunk rounded apex, 2–10 × 1.2–5(–6) mm. Pollen
clear, drying white A. araucana cones 2–12 cm long and 3–22 mm wide 7
7a. Adult leaves subulate or lanceolate 8
7b. Adult leaves (ob)ovate or elliptic-ovate 11
Key to the species of Section Eutacta
8a. Pollen cones very slender, 3–7 mm wide; micro­
1a. Foliage branches with and including adult sporophylls connate. Outer bark on trunks ex-
leaves (12–)15–50 mm wide; branching archi- foliating with thin, curling flakes
tecture according to Rauh’s model 2 A. cunninghamii
1b. Foliage branches with and including adult 8b. Pollen cones 7–20 mm wide; microsporophylls
leaves (2.5–)3–13(–16) mm wide; branching ar- imbricate or divergent. Outer bark on trunks
chitecture mostly according to Massart’s model exfoliating in horizontal strips 9
(in one species Rauh’s model) 5 9a. Pollen cones 2–5 cm long, 7–11 mm wide. Seed
2a. Foliage branches with and including adult cones when mature 7–9 × 5–6 cm. Foliage
leaves 30–50 mm wide; adult leaves 20–32 × branches with and including adult leaves 6–10
9.5–18.5 mm, triangular to linear-triangular mm wide. Endemic to the summit of Mt. Panié
A. muelleri in New Caledonia A. schmidii
2b. Foliage branches with and including adult 9b. Pollen cones 5–12 cm long, 10–20 mm wide.
leaves (12–)15–33 mm wide; adult leaves 8–20 Seed cones when mature 7.5–12 × 6–10 cm. Fo-
(–25) × 4–14 mm, broadly lanceolate, ovate or liage branches with and including adult leaves
nearly triangular 3 4.5–8(–12) mm wide. Occurring elsewhere in
3a. Trees with sparse but long branches; crown New Caledonia 10
candelabra shaped, with tufts of thick foliage 10a. Microsporophylls of pollen cones divergent,
branches at ends; adult leaves broadly lanceo- spreading at nearly 90° from the rachis, ovate,
late, short retained on leading branches 5–8 × 4–5 mm. Adult leaves 4–10 mm long
A. rulei A. nemorosa
3b. Trees with numerous branches; crown colum- 10b. Microsporophylls of pollen cones imbricate,
nar, broadly conical or domed; foliage branches spreading at 45° from the rachis, triangular, 3–4
usually not limited to ends of main branches; × 2–2.5 mm. Adult leaves (3–)4–6 mm long
adult leaves broadly ovate to nearly triangular, A. subulata
long retained on leading branches 4 11a. Crown of mature trees domed to flat-topped.
4a. Adult leaves nearly triangular. Crown colum- Microsporophylls of pollen cones connate, 2.5
nar or broadly conical. Mature seed cones 10–13 × 2 mm A. scopulorum
 11b. Crown of mature trees columnar or narrowly (perhaps originally Amerindian) and many other
conical. Microsporophylls of pollen cones im- variants denoting ecotypes or seed types.
bricate, 3–7(–10) × 2–4 mm 12
12a. Foliage branches with and including adult
Description
leaves very slender, 3–7 mm wide; adult leaves
broadly ovate, 2.5–4.7 × 1.5–2.5 mm A. bernieri Dioecious or less commonly monoecious trees to 50
12b. Foliage branches with and including adult m tall, to 2 m d.b.h.; trunk straight. Bark to 15 cm
leaves slender to more robust, (3–)5–12 mm thick, rough and deeply fissured, exfoliating in
wide; adult leaves elliptic-ovate or ovate, 4–8 × small chips and plates, sometimes exposing smooth
1.8–5(–6) mm 13 patches of reddish inner bark; outer bark grey. Resin
188 13a. Crown of mature trees columnar but open, in exudate reddish. Crown in mature trees domed and
older trees with a flat or candelabra shaped top, finally flat-topped, with branches only at the top of
in young trees pyramidal. Pollen cones small, the tree; branching according to Rauh’s model. Pri-
3–3.5 cm long and 9–10 mm wide mary branches in pseudo-whorls of 4–8, up to 5 m
A. humboldtensis long, spreading or assurgent, mostly with second to
13b. Crown of mature trees narrowly columnar third order branches, caducous. Adventitious foli-
or narrowly conical, with a narrowing top, in age branches uncommon, usually associated with
young trees (narrowly) conical. Pollen cones damage. Foliage branchlets spreading or ascend-
(2.5–)4–7(–10) cm long and 10–22 mm wide 14 ing (sometimes more or less pendent) on primary
14a. Crown of mature trees narrowly columnar. Pol- branches, forming large tufts of foliage, of unequal
len cones 13–22 mm wide; microsporophylls length up to 50 cm, up to 40 mm wide (including
oblong-triangular, 5–7(–10) × 2.5–3(–4) mm leaves), but of unequal width from base to apex de-
A. columnaris pending on size and spread of leaves, more or less
14b. Crown of mature trees narrowly conical. Pollen flexible. Adult leaves imbricate or variously spread-
cones 10–15 mm wide; microsporophylls rhom- ing (40–90°), decurrent, ovate to lanceolate, 1.5–5 cm
bic, 3–4 × 4 mm A. heterophylla × 3–20 mm, varying much in length and width often
on a single branch, flat or slightly concave adaxially,
sometimes weakly keeled abaxially, smooth or stri-
Araucaria angustifolia (Bertol.) Kuntze, Rev. Gen. ate, tapering to a pungent apex; margins entire. Sto-
Pl. 3: 375. 1898. Columbea angustifolia Bertol., mata in 40–50 indistinct rows on the abaxial surface
Opusc. Sci. 3: 411. 1819. Type: Brazil: Rio de Janeiro, and slightly fewer but more conspicuous rows on the
Corcovado, [“In Brasilia legit G. Raddi”], G. Raddi adaxial surface. Pollen cones axillary at end of foli-
s.n. (holotype BOLO). Pl. 6 age branchlets, solitary or in small clusters, initially
erect, cylindrical and green, becoming pendulous,
Araucaria angustifolia (Bertol.) Kuntze var. depen­ elongated, curved and pinkish brown at anthesis,
dens J. R. de Mattos, Loefgrenia 21: [2]. 1967. 8–15 cm × 15–30 mm. ­Microsporophylls connate,
Araucaria angustifolia (Bertol.) Kuntze var. vinacea spreading at ca. 90° from a stout rachis, apically free
J. R. de Mattos, Loefgrenia 111: 1. 1997. at anthesis; stalk thin and weak, linear, 3–5 mm long;
lamina cuneate, rhombic in outline, 4–5 × 3 mm,
lateral margins entire; apex curved; abaxial sur-
Etymology
face smooth. Pollen sacs 8–12 (–15), linear, 6–8 mm
The species epithet alludes to the (often) narrower long. Seed cones axillary on stout foliage branches
leaves than those of A. araucana, the only other spe- which widen below the cone, usually 3–5 together in
cies native to South America. whorls, erect. Immature cones ovoid, 4–6 cm long,
densely covered with recurved bract tips, green,
maturing in 20–22 months. Mature cones chestnut
Vernacular names
brown, ovoid-globose to globose, usually slight-
Parana pine, Candelabra tree; pinheiro-do-Paraná, ly longer than wide, (8–)12–20 × 8–18 cm (green
also recorded are: pinhao, pinheiro, pinho-nacional weight 3–4 kg). Bracts ovoid-oblong to cuneate,
(Portuguese); pino Paraná (Spanish); curiy or kuriy more or less angular, ca. 5 × 2 cm including vestigial
wings if present, distally thickened to a more or less
 189

5 4 3

Pl ate 6 . Araucaria angustifolia. 1. Habit of tree. 2. Foliage branch with pollen cone. 3. Leaf. 4. Pollen
cone. 5. Microsporophyll with pollen sacs. 6. Seed cones, one with top removed to show seeds.
 rhombic, transversely keeled apo­physis, ending in s­ ummer rains in much of its range, but with more
a caudate, slightly curved, 8–10 mm long tip. Seed winter rains in the south, to a total of 1400–2500
scale not wider than the seed; ligule fragile, 1–1.5 mm per annum. The prevalent soil type in the Arau-
mm long, sometimes absent. Seeds ovoid-­oblong, caria forests is a well-drained, iron-rich red-yellow
ca. 4 × 1.5 cm, not flattened, striated longitudinally podzol with a low pH value.
or smooth, ripening lustrous red-brown (sometimes
striped darker red, or with white or pale apex, or oc-
Conservation
casionally whole seed very pale).
At the beginning of the 20th century accord-
ing to estimates the area of occupancy (AOO) of
Distribution
190 A. angusti­folia was around 20 million ha. Based
Argentina, S Brazil, Paraguay. The range of this spe- on LANDSAT-II imagery, it was calculated that by
cies before European settlement was extensive and 1982 only 565,419 ha remained and this was still be-
predominantly located in southern Brazil, with ex- ing logged at a rate of almost 80,000 ha per year
tensions into eastern Paraguay and the Argentinian (Gantzel, 1982). Further statistics are given by Koch
Misiones region between the Paraná and Uruguay & ­Celeste Correa (2002). AOO was ca. 200,000 km²
Rivers. To the NE its range is broken up into more on arrival of European colonists in the 19th centu-
scattered occurrences. ry; it is now down to 4000 km². The greatest losses
TDWG codes: 84 BZL-MG BZL-SP BZS-PR BZS-RS occurred between 1915–1960, by the latter year the
BZS-SC 85 AGE-MI PAR area was down to 20,000 km², but most of that has
now also gone. Based on these figures, the species
is Critically Endangered under criterion A1cd even
Ecology
though exploitation rates may have been drastically
This species is a dominant and/or emergent tree in reduced as logging of natural stands is now prohib-
temperate to subtropical forests or forest enclaves ited by law. Even that is doubtful, as illegal logging is
(capâo) amid extensive grassland (campo limpo), or reported to continue, even in nature reserves, while
sometimes in small isolated groups of trees. Altitu- many remaining stands are in private hands, where
dinal range: (300–)500–1800 m a.s.l. Its most exten- the owners consider the trees their disposable prop-
sive and more or less continuous occurrence (before erty. Plantation of the species has been substantial,
large scale deforestation through logging) was on but that does not make up for the loss of ‘old growth’
the Planaltos, a more or less level plain at medium under IUCN criteria. Economic factors have caused
altitudes between 500 m and 1000 m. Tree ferns the replacement of Araucaria by faster growing
(­Alsophila, Dicksonia) and also many deciduous as plantations of Pinus and Eucalyptus in many areas,
well as evergreen smaller trees and shrubs form one while only few landowners are inclined to substan-
or more understoreys. Most common among decid- tially replant with A. angustifolia, preferring exotics
uous trees is Cedrela fissilis; other angiosperm trees for that purpose.
are Cryptocarya aschersoniana, Drimys brasiliensis IUCN: CR (A1cd)
and the genera Leandra, Miconia, Ocotea, Nectandra
and Tibouchina, as well as the tall palm Arecastrum
Uses
romanzoffianum. Podo­carpus lambertii and, locally,
P. sellowii are two other conifers growing under or This species remains one of the most valuable tim-
with A. angustifolia, but they are less common and ber trees of Brazil, despite its very substantial re-
indicate undisturbed forest. The climate varies with duction in total volume of standing timber due to
latitude and altitude; on the Plan­altos at around sustained over-exploitation. It is known as Paraná
1000 m frost is a common occurrence in winter pine in the trade. It is essentially the only native
with minima to –10 °C while snowfall, although conifer of large size to produce long, straight boles
rare, can occur in most of the range of the species. of excellent sawing quality. Its wood is soft and light
According to Golte (1993) the northern limit of and large, straight trees have little or no knots from
natural ­occurrence of A. angusti­folia coincides with branches. Paraná pine ‘knot’ wood, unsuitable for
the zone where tropical winter dry seasons begin to anything else, was burnt in (among other things)
have influence upon the ­vegetation. Rains are ­mostly steam locomotives. The uses of its wood have been
highly ­varied over the years of exploitation, from Rauh’s model. Primary first order branches of ma-
construction timber to matches and toothpicks and ture trees in pseudo-whorls of 8–10 or sometimes
from pulp for paper making to plywood and musical more, up to 5 m long, spreading or assurgent, mostly
instruments. The knot powder is used in Brazil in with second to third order branches, caducous. Ad-
flexographic inks, plastic laminates, furniture var- ventitious foliage branches common, but usually
nishes, and as a partial substitute for phenolic resins. associated with damage, showing a tendency to pro-
In rural areas, houses and farmsteads were built of it. duce secondary crowns. Foliage branchlets spread-
Plantations are generally not favoured by the timber ing or ascending on primary branches, of unequal
trade as growth of Araucaria is much slower than length up to 1 m or more, 6–9 cm wide (including
that of introduced eucalypts and pines. Despite this, leaves), of regular width from base to apex, rigid,
there are now some plantations of this species in the lower foliage branches sometimes sub-pendulous. 191
region aimed at harvesting trees in due course. Its Adult leaves variously spreading (45–90°), with a
seeds are edible and are eaten by the Brazilian Am- decurrent, reflexed base merging with bark, ovate to
erindians in a similar manner to bunya nuts (A. bid- lanceolate, 2.5–6 × 1.5–3 cm, varying little in length
willii) in Australia and the seeds of A. araucana in and width on a single branch, flat or slightly con-
Chile and Argentina. The tree is also sacred to sev- cave adaxially, ca. 1 mm thick, rigid and coriaceous;
eral of the Amerindian tribes, such as the Kaingang margins entire; surface smooth or striate, abaxially
people of the Paraná River in S Brazil who regard it keeled and gradually or more abruptly tapering to a
as a masculine symbol. Although still uncommon, it pungent apex. Stomata in irregular and inter­mittent
is seen more often recently in arboreta and large gar- lines from leaf base to apex, rows more numerous
dens in milder parts of Europe and North America on the abaxial surface than on the adaxial. Pollen
(Grimshaw & Bayton, 2009: 152). cones axillary, solitary or more often in small clus-
ters, initially erect, stoutly cylindrical or slightly
fusiform, becoming elongated and curved down at
Araucaria araucana (Molina) K. Koch, Dendrol. anthesis, 8–15 cm × 40–50 mm. Microsporophylls
2 (2): 206. 1873. Pinus araucana Molina, Sag. Stor. imbricate, spreading at ca. 60° from a 10 mm thick
Nat. Chili: 182. 1782. Type not designated. Fig. 41, rachis; stalk thin and weak, ca. 10 mm long, straight
42, 43 or slightly curved; lamina 15–20 mm long, 4–5 mm
wide at the thickened and curved base, lateral mar-
gins denticulate; apex acuminate, recurved; abax-
Etymology
ial surface slightly rugose. Pollen sacs 10–15, linear,
The species epithet is derived from the Araucanos, ca. 10 mm long, tightly packed. Seed cones axillary
a group of Amerindian tribes of the Araucana lin- on very short foliage branches which widen below
guistic family. cone, usually solitary, erect. Immature cones ovoid,
6–8 cm long, densely covered with recurved bract
tips, green, maturing in 22–24 months. Mature
Vernacular names
cones dull brown, subglobose to globose, 15–20 cm
Monkey Puzzle, Chilean pine, Araucaria; chihuén, diam. (green weight 3–4 kg). Bracts ovoid-oblong
also recorded are pehuén, araucaria de Neuquén, to cuneate, more or less angular, 4–6 × 1.5–2 cm in-
pino de Neuquén, pino hachado, pino solo (Spanish, cluding vestigial wings if present, distally thickened
with some influence from Amerindian languages). to a more or less triangular, transversely keeled apo­
physis, ending in a long-caudate, curved, 2–2.5 cm
long tip. Seed scales not wider than the seed; ligule
Description
fragile, ca. 2 mm long, sometimes absent. Seeds ob-
Dioecious or more rarely monoecious trees to 50 long-conical, 3.5–5 × 1–1.5 cm, not flattened, smooth,
m tall, to 2.5 m d.b.h.; trunk straight. Bark to 15 cm matt tawny brown when ripe.
thick, rough and deeply fissured, exfoliating in small
chips and plates; inner bark brown; outer bark grey
Taxonomic notes
or grey-brown. Crown in mature trees variously
domed or more or less flat-topped, with branches Araucaria araucana shows surprisingly little mor-
only at the top of the tree; branching according to phological diversity among populations in Chile as
 well as between cultivated trees elsewhere. Due to fagus at such sites. It is a typical example of a tree
its size and harsh nature the foliage has rarely been species well adapted to episodic, large scale distur-
­collected for herbaria and most botanists know it bance events.
primarily from the numerous planted trees, espe-
cially common in NW Europe. Original material
Conservation
does not exist and a neotype would therefore be nec-
essary to fix the accepted name. Its nearest relative In the Andes, the populations are scattered and frag-
species, also from South America, is A. angustifolia. mented and its area of occupancy (AOO) may not
have exceeded 4000 km2 in historical times (Golte,
1993). The limited AOO of this species in the wild,
Distribution
192 combined with the scattered nature of its remaining
S Argentina: Neuquén; S Chile: Biobio, La Arau- populations, put it in the category Vulnerable under
cania, Los Lagos. Mainly in S Chile, but part of the IUCN criteria. Between the 1920s and 1970s, the area
­Andean distribution is across the border in Argen- covered by the species was halved as a result of log-
tina. ging and fires. While (illegal) logging has played a
TDWG codes: 85 AGS-NE CLC-BI CLC-LA CLS-LL part in its historical decline, most stands are now on
protected lands. However, these are largely situated
in proximity of active volcanos with their inherent
Ecology
periodic eruptions causing forest destruction. Fires
To the north, A. araucana is limited by a summer- have in recent years caused substantial damage to
dry Mediterranean type climate, to the south by several subpopulations. The limited occurrence in
an increasingly cooler and wetter oceanic temper- the Coastal Cordillera is only partly protected and
ate ­climate; its natural distribution therefore marks threatened by habitat loss and conversion to planta-
a narrow transitional zone between two floristic tion forestry. Sufficient recruitment through natural
realms: neotropic and antarctic (Golte, 1993). Espe- seed dispersal and regeneration to replace such loss-
cially in the Andes, its distribution is closely associ- es may be hampered by a number of factors which
ated with volcanic activity, primarily with deposits have increased their impact through human inter-
of tephra and scoria; the coastal mountains are com- ference, e.g. the harvesting of seeds for food mar-
posed of granitic or metamorphic rocks. Altitudinal kets. The species is listed on Appendix I of CITES
range: 600–1800 m a.s.l. Both coastal mountains and as a Natural Monument in Chile, which gives
and Andes receive high amounts (2000–3000 mm) it legal protection against logging. However, during
of annual precipitation within the altitudinal zone the 2001/02 fire season, catastrophic fires caused by
of the araucarias, but also experience dry and hot lightning destroyed much of the remaining available
spells in summer. Araucaria araucana builds typical habitat including 60% of Tolhuaca National Park.
stands either pure or at lower altitudes mixed with Additional protection through active management
Valdivian rain forest angiosperm trees especially of some of these factors would be necessary to en-
Nothofagus dombeyi; towards the tree line in the sure the existence of healthy and sufficiently large
Andes only shrubs, especially Nothofagus pumilio, populations throughout the natural area of extent of
or small trees like N. antarctica and Drimys win- A. araucana, and especially in the disjunct coastal
teri, grow under the conifers. Easternmost popu- subpopulations, one of which is genetically distinct
lations in Argentina occur in areas with much less from all the others.
precipitation (1000–1500 mm) and here the conifer IUCN: VU [A2c; B1ab(ii,iii,v)]
Austrocedrus chilensis can be found associated with
A. araucana. Araucarias are capable of colonizing
Uses
fresh falls of tephra or scoria after volcanic eruptions
before other tree species and maintain dominance, Before logging was outlawed, A. araucana was one
slowly increasing stand density and so shading out of the most desirable sources of sawn timber in
competitors. Trees killed off above ground can re- Chile. The edible seeds are traditionally harvested by
generate from epicormic buds at the base or even the Mapuche Indians, particularly a mountain tribe
from shallow roots. Resistance to low intensity fires called the Pehuenche. They are eaten raw, boiled, or
also helps to maintain its dominance over Notho­ toasted in the ashes of a fire or in a pot or stove, or
ground into flour that can be used to make bread towards the end, 3–7 mm wide (including leaves),
or as a condiment in soups; they are also made into remaining of equal width almost throughout, flex-
a fermented drink known as chavid. As well as be- ible. On mature trees leaves of two kinds: adult
ing eaten during the collection season, they are pre- leaves on primary branches and semi-juvenile leaves
served for year-round use, either by hydration in a on adventitious branches. Adult leaves on ultimate
pit of cold water or by dehydration in the sun or over branchlets broadly ovate, 2.5–4.7 × 1.5–2.5 mm,
the family fire. Piñones are now increasingly traded 3-edged and incurved with acute or obtuse apex,
(on rather unfavourable terms) in local and regional smooth, prominently keeled abaxially; leaves sub-
markets for yerba maté, sugar and other foodstuffs tending seed cones 8–10 × 6–8 mm, with a rostrate
as well as for cash. Araucaria araucana was intro- apex, much larger than normal foliage leaves. Sto-
duced into Europe soon after its discovery in the mata on the abaxial side of adult leaves restricted to 193
late 18th century, it proved to be winter hardy in most the proximal part of the leaf; on the adaxial side nu-
countries and in the 19th century became a fashion- merous in rows from just above base to apex. Semi-
able tree for large gardens, parks, and stately avenues juvenile leaves divergent, oblong, curved, 4–6 × 1–2
to country houses. Trade for horticultural purposes mm, widest at base, three-faced, bilaterally flattened;
is now entirely based on plants raised in cultivation apex obtuse. Pollen cones terminal, cylindrical, ini-
due to an export ban under the CITES convention. tially erect and straight but pendulous and curved
at anthesis, light brown with glaucous bloom, (2.5–
)3.5–4.5(–10) cm × 8.5–10 mm. Microsporophylls
Araucaria bernieri J. T. Buchholz, Bull. Mus. Hist. spreading at 45–90° from a stout rachis; stalk thin
Nat. (Paris), sér. 2, 21: 280. 1949. Type: New Cale­ and weak, linear, 2 mm long; lamina ovate, 3–4 ×
donia: Grande Terre, Province Sud, Rivière des 2–2.5 mm, curved at base; apex acute or pungent.
Pirogues, near Lucien’s sawmill, J. T. Buchholz 1562 Pollen sacs 4–6, linear, 2–2.5 mm long. Seed cones
(holotype ILL). Fig. 44 terminal on short, stout foliage branches which
widen below the cone, usually solitary, sometimes
2–3 together, erect, subglobose, often widest above
Etymology
the middle, 8–10 × 6.5–8 cm. Bracts flabellate, 2.5–3
Julien Bernier was the first Conservator of the New × 2–2.5 cm including rounded, thin membranous
Caledonian Museum, Nouméa; his son (another wings, distally thickened but thin-edged, ending in
J. Bernier) collected a fragment of A. bernieri near a caudate, slightly curved, 6–8 mm long bract tip.
Pic Busse. Seed scales not wider than the seed; ligule broadly
triangular, fragile, 1.5–2 mm long. Seeds obovoid,
15–20 × 6–8 mm, transversely flattened, striated lon-
Vernacular names
gitudinally or smooth, light brown when ripe.
No common names have been recorded for this spe-
cies.
Distribution
New Caledonia: Grande Terre, restricted to the ex-
Description
treme south and on heights along the SE coast.
Monoecious trees to 50 m tall, to 1 m d.b.h.; trunk TDWG codes: 60 NWC
straight or curved. Bark exfoliating in horizon-
tal strips; inner bark dark reddish brown; outer
Ecology
bark grey or brown. Crown columnar but open,
branched along greater part of trunk, often widest This species is commonly found in angiosperm
in top; branching according to Massart’s model. Pri- ever­green rainforest on slopes of lower mountains,
mary branches followed by infrequent adventitious with an altitudinal range of (1–)100–500(–800) m
branching in the lower ¾ of the crown; first order a.s.l. It does not form dense stands but rather scat-
branches in pseudo-whorls of 5–7(–10), 0.5–2.5 m tered individual trees that emerge well above the
long, spreading or assurgent, caducous. Foliage closed canopy of the forest, sometimes in associa-
branchlets of more or less equal length ((8–)19–35 tion with A. subulata. In some localities A. bernieri
cm) on either side of the axis branch and shortening has been observed to follow stream beds in ravines.
 A few collections were made from localities near sea
Description
level in the extreme south of the island, usually near
streams lined with closed forest patches. Araucaria Monoecious trees to 45 m tall, to 1.75 m d.b.h., with
bernieri seems nearly everywhere to be limited to straight trunk. Bark 7.5–10(–30) cm thick, rough
soils derived from ultramafic rock (peridotite and and scaly, exfoliating in thin plates and scales; in-
serpentine) with a high content of heavy metals ner bark orange to reddish; outer bark grey to black.
such as iron, manganese, and nickel. It is likely that Resin exudate clear, weathering white to grey. Crown
the forest canopy and resulting leaf litter prevent in mature trees domed and open especially in the
de­hydration of these permeable soils, which can lower part, with primary branches restricted to
quickly occur despite very high annual rainfall in upper part of tree; branching according to Rauh’s
194 this region (2000–3000 mm). Where the forest has model. Primary first order branches of mature trees
been disturbed A. bernieri may survive in stream in pseudo-whorls of 10–16, up to 7 m long, spread-
beds with permanent (underground) water. ing, down-curved or assurgent, caducous. Adventi-
tious foliage branches common. Foliage branchlets
spreading in all directions or pendent on ends of
Conservation
primary branches, forming large tufts of foliage, of
Mining is a potential threat to A. bernieri because unequal length up to 60 cm, up to 4 cm wide (in-
it is virtually restricted to ultramafic rock types cluding leaves), of unequal width from base to apex,
with high metallic content. Deforestation by cut- flexible or more rigid. Leaves present on first to third
ting and burning has been extensive within its range order branches, retained on (pen)ultimate branches
and many populations are in isolated remnants of which are caducous in mature trees. Adult leaves
rain forest on slopes and in hollows and ravines imbricate or variously spreading (radially to disti-
of mountains. Some of these have been observed chously), dimorphic: on orthotropic shoots narrowly
to have been recently affected by fires. The species triangular to acicular, on plagiotropic shoots trian-
is still fairly common in the south and occurs in a gular to lanceolate, 10–50 × 3–15 mm, varying much
number of protected forest areas, e.g. Montagne des in length and width often on a single foliage branch,
Sources, Pic du Grand Kaori and Rivière Bleue. nearly flat in long leaves to navicular in short leaves,
IUCN: VU (C1) smooth or striate or faintly keeled, tapering to an
acuminate apex, pungent or not. Stomata in 35–80
irregular and intermittent rows from base to apex on
Uses
each surface. Pollen cones axillary, in small clusters
The wood is of high quality and has been locally ex- of 2–6, narrowly cylindrical, initially erect, becoming
ploited. elongated, curved to pendulous at anthesis, 6–15 cm
× 8–15 mm. Microsporophylls connate, spreading at
ca. 90° from a thin, fragile rachis; stalk thin and weak,
Araucaria bidwillii Hook., London J. Bot. 2: 3 mm long; lamina curved, the exposed part rhombic
503, t. 18, 19. 1843. Marywildea bidwillii (Hook.) in outline, 3 × 2 mm. Pollen sacs 8–10, linear, ca. 3 mm
A. V. Bobrov & Melikyan, Komarovia 4: 58. 2006. long. Seed cones axillary (appearing sub-terminal)
Type: Australia: Queensland, Wide Bay District, on a very short, stout and leafy peduncle, usually soli-
Mt. Brisbane Range, [“Wide Bay” near Gympie], tary, erect. Mature cones green, ovoid-globose to glo-
J. C. Bidwill s.n. (lectotype K). Fig. 45, 46 bose, usually slightly longer than wide, 20–30(–35) ×
20–25 cm (green weight typically 4–5 kg but up to 10
kg), often falling when still intact. Bracts broadly ob-
Etymology
trullate to cuneate, 10–12 × 10–15 cm, lateral sections
This species was named after the collector, John thin, distal part thickened to a more or less rhombic,
Carne Bidwill (1815–1853). sharply keeled apophysis, ending in a slightly curved,
10–15 mm long, caducous tip. Seed scales not wider
than the seed; ligule thick, c. 15 × 10 mm. Seeds deeply
Vernacular names
sunken in the seed scale, ovoid-cuneate, 4–5 × 2.5–3
Bunya pine (a corruption of the Aboriginal name cm, slightly flattened, smooth, ripening lustrous
bon-yi). brown and breaking free from cone scale.
 Distribution Uses
Australia: Queensland. Two disjunct areas ca. 1000 Bunya pine produces excellent seeds for human
km apart: in S Queensland in the Blackall Ranges, consumption. The species is sacred to the Austral-
Bunya Mts., Brisbane River, Mary River Valley and ian Aborigines; its importance to Aboriginal culture
Yarraman/Blackbutt area; in N Queensland at Can- was celebrated in a series of articles published in
nabullen Falls and Mt. Lewis. a special issue of the Queensland Review in 2002.
TDWG codes: 50 QLD-QU The Aborigines used to meet in years of good seed
crops (cones mature in three years) in large num-
bers for harvest and festivities. Each clan or tribe
Ecology
had jurisdiction over certain trees and their crop 195
Araucaria bidwillii is a large, emergent tree in (sub) of seeds. Their traditional harvesting rights have
tropical rainforest (‘notophyll vine forest’) on basal- not been upheld; their last great gathering for the
tic or other igneous substrates; sometimes associated bon-yi harvest was held in 1902. Interest in aborigi-
with A. cunninghamii, raising its domed crowns well nal foods, ‘bush tucker’ as it is popularly called in
above a canopy of evergreen angiosperm trees. Its Australia, is growing and the seeds of A. bidwillii are
altitudinal range is 150–1000 m a.s.l. Araucaria bid- again on the menu, now also of white Australians.
willii does not form pure stands, but is always mixed The wood is light and easily worked, and used for
to a greater or lesser extent with numerous species flooring, plywood and cabinet-making. In the past
of angiosperm trees mostly belonging to tropical these trees were logged by European settlers, but the
families. Dominant among these are Argyrodendron last sawmill closed in 1945 and most of the remain-
actinophyllum, Drypetes australasica, Ficus macro- ing trees are protected in the Bunya Mountains Na-
phylla, and F. obliqua; also common are the palm tional Park. This species has been widely introduced
Archontophoenix cunninghamiana and tree ferns. as an ornamental in New Zealand, India, Portugal
The forest is divided into larger and smaller woods and other countries.
by ‘balds’, coarse grasslands or open savannas which
form sharp boundaries with the forest patches. An-
nual precipitation is 1100–1400 mm in the southern Araucaria biramulata J. T. Buchholz, Bull. Mus.
area, with a dry season from April/May to Septem- Hist. Nat. (Paris), sér. 2, 21: 279. 1949. Type: New
ber; the northern populations receive a more evenly Caledonia: Grande Terre, Province Sud, Forêt du
distributed annual precipitation of 1500–2000 mm. Mois de Mai (on ridge between rivers Bleue and
Blanc), J. T. Buchholz 1691 (holotype ILL).
Conservation
Etymology
Beginning in the early decades of European settle-
ment and in the Bunya Mountains continuing until The species epithet “…refers to the characteristic
the end of the Second World War in 1945, logging twice-branched character vegetative twigs” (Buch-
must have reduced the original area of occupancy holz, op. cit., p. 280).
of this species, especially in the Blackall Range and
headwaters of the Brisbane River. According to Golte
Vernacular names
(1993), no detailed map of the distribution, past or
present, of A. bidwillii exists in the literature, a situ- None are recorded.
ation that frustrates attempts to evaluate the IUCN
Red List status of this species. Herbarium specimens
Description
that could give clues of past occurrence are few, due
to the difficulty of collecting them. Nearly all re- Monoecious trees to 30 m tall, to 0.8 m d.b.h.; trunk
maining stands are now on protected land and log- straight. Bark to 2 cm thick, exfoliating in horizontal
ging has virtually ceased. strips; inner bark deep wine red; outer bark grey with
IUCN: LC some brown. Resin exudate white or creamy, drying
 opaque and powdery. Crown in mature trees coni-
Ecology
cal or columnar but open, branched along greater
part of trunk; branching according to Rauh’s model. This is one of the New Caledonian species that is
Adventitious branching absent or infrequent. Pri- restricted to peridotite and serpentine. Altitudinal
mary first order branches of mature trees in pseudo- range: 190–1150 m a.s.l. It forms thickets and is often
whorls of 4–5, 1–2.5 m long, ascending, sometimes seen growing on steep rock faces and cliffs, leaning
with second order branches, caducous. Foliage over instead of upright. It also occurs in humid val-
branchlets strongly assurgent on primary branch- ley bottom forest and in more open ‘maquis minier’,
es and confined to distal parts, of unequal length usually on slopes. It is sometimes associated with
(10–37 cm), shortening towards end of branchlet, Agathis lanceolata.
196 (5–)8–13(–16) mm wide (including leaves), variable
in width from base to apex with changing size and
Conservation
spreading of leaves, flexible, often branching again.
In mature trees leaves of two kinds: adult and semi- This species has probably been seriously reduced in
juvenile. Adult leaves on (pen)ultimate branchlets its occurrence on some mountains that are inten-
ovate or broadly lanceolate, (4–)5–10 × 3–6 mm, ± sively mined for nickel, especially in Province Nord.
naviculate with incurved, subacute non-pungent On Mt. Kaala and the Massif de Kopéto the species
apex, coriaceous, smooth, lustrous green, rather has not been found since 1980; from the first moun-
weakly keeled abaxially; leaves subtending seed tain it has almost certainly gone (pers. obs. No-
cones 10–16 × 2.5–4 mm, with a rostrate apex. Sto- vember 2005). Since the number of mature trees is
mata on the abaxial surface of adult leaves mostly in estimated to be fewer than 10,000 this species meets
the proximal half of leaf; on the adaxial surface 10– the criteria for Vulnerable.
14 rows from base to apex on each side of the mid- IUCN: VU [B1ab(iii); C2a(i)]
vein. Semi-juvenile leaves on adventitious branches
spreading more widely than adult leaves on primary
Uses
branches, lanceolate, 4–8 × 2–4 mm, straight in low-
er 2/3 but distally incurved, keeled abaxially; apex No uses have been recorded of this species.
subacute and shortly mucronate. Pollen cones ter-
minal on the ultimate foliage branchlets, cylindri-
cal, initially erect, pendulous and curved at anthesis, Araucaria columnaris (J. R. Forst.) Hook., Bot.
6–7 cm × 15–20 mm. Microsporophylls imbricate, Mag. 78: sub t. 4635. 1852. Cupressus columnaris
spreading at 45° from a stout rachis, apically free at J. R. Forst., Fl. Ins. Austr.: 67. 1786. Type: New Cale­
anthesis; stalk thin and weak, 3–4 mm long; lamina donia: Province Sud, Île des Pins, W. Anderson s.n.
triangular-­apiculate, 4–5 × 3–4 mm; abaxial surface (holotype BM). Fig. 47, 48
rugose. Pollen sacs 7–8, ca. 3.5 mm long. Seed cones
terminal on short, stout foliage branches which
Etymology
widen below cone, usually solitary, sometimes 2–3
together, erect, subglobose, 9–11 × 8–10 cm, with a The species epithet refers to the columnar habit of
truncate apex. Bracts flabellate, 2.5–3 × 2–2.5 cm in- this species.
cluding rounded, thin membranous wings, distally
thickened and prominently keeled, ending in a cus-
Vernacular names
pidate, curved, 8 × 2.5 mm tip. Seed scales not wider
than the seed; ligule fragile, 2–2.5 mm long. Seeds Cook’s pine, New Caledonia pine; pin colonnaire
obovoid-oblong, 20–25 × 8–10 mm, transversely (French).
flattened, smooth, light brown when ripe.
Description
Distribution
Monoecious trees to 50(–60) m tall, to 1–1.5 m d.b.h.;
New Caledonia: Grande Terre. trunk straight or curved. Bark to 3 cm thick, exfo-
TDWG codes: 60 NWC liating in horizontal strips, forming small, curled
 flakes; inner bark dark reddish brown; outer bark
Ecology
light grey. Resin exudate yellow to orange, drying
white or brown. Crown in mature trees narrowly Araucaria columnaris occurs on coral reefs and
columnar, branched along greater part of trunk; coastal headlands in proximity of the sea in natural
branching according to Massart’s model. Primary habitats, and is an emergent. It is widely planted else-
branches followed by frequent adventitious branch- where in New Caledonia, especially on coastal head-
ing. Primary first order branches of mature trees in lands and islands and in some valleys of the interior.
pseudo-whorls of 5–7(–10), 1–2(–3) m long, spread- It occurs in the wild mostly, but not exclusively, on
ing or assurgent, caducous. Foliage branchlets in calcareous substrate, often on coral. Occasionally, as
mature trees confined to distal parts of primary at Port Boisé, it is found on basalt or serpentine sub-
branches (and on adventitious branches), of vari- strates. In the southern part of Grande Terre it is re- 197
able length (2–40 cm), 5–10 mm wide (including stricted to off-shore reefs and islets and some coastal
leaves), more or less terete, flexible, caducous. In slopes on ultramafic rock or soil, from just above
mature trees leaves of two kinds: adult leaves on pri- high tide to ca. 50 m a.s.l.; on the Loyalty Islands
mary branches and semi-juvenile leaves on adventi- and the Île des Pins it grows on low lying calcare-
tious branches. Adult leaves on ultimate branchlets ous rock derived from coral reefs. The tall trees are
­elliptic-ovate to obovate-elliptic with the shortest emergent high above a canopy of sclerophyll scrub
leaves ovate-lanceolate, 4–8 × 1.8–5 mm, rhythmi- (‘maquis minier’) or dry forest with e.g. Pandanus
cally varying in shape and size along a branch, more bernardii and grow as scattered individuals either
or less navicular with incurved, acute or obtuse apex, more or less solitary or in more or less dense ‘groves’.
smooth, keeled abaxially; leaves subtending seed Where the forest is less dense dominant associates
cones more or less triangular, 8–14 × 4–8 mm. Sto- include Argusia argentea, Cycas seemannii, Scaevola
mata on adult leaves of ultimate branchlets abaxially taccada, Pandanus tectorius, Premna integrifolia,
confined to the proximal part of the leaf, on adax- Boerhavia repens and Pisonia grandis. On beaches,
ial surface in 30–33 rows. Semi-juvenile leaves di- A. columnaris grows in association with Acacia sim-
vergent, erecto-patent, narrowly oblong-lanceolate, plicifolia, Calophyllum inophyllum and Casuarina
falcate, on branchlets 5.5–11 × 0.6–1.5 mm, ending equisetifolia. On the ultrabasic rocks at Port Boisé,
in a short blunt tip. Pollen cones terminal, solitary, Araucaria columnaris is associated with the local en-
­conical-cylindrical, initially erect, pendulous and demic A. nemorosa.
curved at anthesis, (2.5–)4–7(–10) cm × 13–22 mm,
lax. Microsporophylls spreading at 45° from a stout
Conservation
rachis; stalk thin and weak, 3–5 mm long; lamina
oblong-triangular, 5–7(–10) × 2.5–3(–4) mm; apex Although it is not always possible to determine with
acute and ± cuspidate. Pollen sacs 10–12 or more, certainty the distinction between natural stands and
linear, 4–6 mm long. Seed cones terminal on short, trees planted in the past within the assumed natu-
stout foliage branches, usually solitary, sometimes ral range of this species, it is obvious that extensive
2–3 together, erect, subglobose, 8–10 × 7–8.5 cm natural stands still occur. There is no longer any
when mature. Bracts flabellate, 2.5–4 × 3–4.5 cm in- timber exploitation of these stands anywhere and
cluding rounded, thin membranous wings, distally due to their close proximity to the coast and pre-
thickened but thin-edged, ending in a caudate, up- dominantly calcareous substrate nickel mining is
ward curved, 8–10 mm long tip. Seed scales not wid- not an issue. Trees may be thrown in high storms
er than seed; ligule triangular, 2–3 mm long. Seeds (typhoons), but regeneration is abundant in many
obovoid, 18–30 × 6–12 mm, more or less transversely places. The trees also enjoy traditional protection
flattened, striated longitudinally or smooth, light from the Polynesian Kanaks and it is generally pro-
brown when ripe. hibited to fell them.
IUCN: LC
Distribution
Uses
New Caledonia: Grande Terre (Province Sud), Île
des Pins, Loyalty Islands. The wood is of high quality and has been locally ex-
TDWG codes: 60 NWC ploited in the past. Araucaria columnaris is sacred
 to the Kanak Polynesian people of New Caledonia, minutely denticulate; leaves subtending seed cones
who regard it as a male symbol and have planted it larger than lower leaves, less curved, with a widened
widely in their settlements. This species is also in- base. Stomata abaxially in 6–7 rows on each side
troduced as an ornamental tree in other islands in of the keel; adaxially in numerous rows extending
the SW Pacific and in India, where it is a popular from base to apex. Semi-­juvenile leaves divergent,
landscape tree in the foothills of the Himalaya and loosely adpressed forwards, lanceolate-elliptic to
in the Western Ghats. ovate-lanceolate; apex subacute and rostrate, shortly
pungent. Pollen cones terminal on ultimate foliage
branchlets, solitary, cylindrical, initially erect, pen-
Araucaria cunninghamii Aiton ex A. Cunn., in dulous and curved at anthesis, 2–5.5(–10) cm × 3–7
198 Lambert, Descr. Pinus, ed. 3, 2, p. s.n. inter 144–145. mm. Micro­sporophylls connate, spreading at ca. 90°
1832. from a slender rachis; stalk thin and weak, 2.5 mm
long; lamina peltate, rhombic or rounded in outline,
2 × 1.5–2 mm; apex acute. Pollen sacs 4–5, pendent,
Etymology
oblong, 2 × 0.6 mm. Seed cones terminal on stout
This species was named after the plant collector Al- foliage branches, usually solitary, erect, ovoid to
lan Cunningham (1791–1839). broadly ovoid or ellipsoid, 5.8–12 × (3.7–)4.5–8 cm.
Bracts flabellate, 2–3 × 2.5–3.5 cm including mem-
branous wings, distally thickened to a transversely
Vernacular names
keeled apophysis, ending in an acuminate, curved,
Hoop pine; several Australian Aboriginal names 5–8 mm long tip. Seed scales not wider than seed,
and local New Guinea names have been recorded in abruptly terminating in a 2–3 mm long ligule. Seeds
the literature about this species. oblong, more or less almond-shaped, 15–20 × 5–8
mm, flattened, brown when ripe.
Description
Distribution
Monoecious trees to 66 m tall, to 2 m d.b.h.; trunk
straight. Bark to 10 cm thick, exfoliating with thin, New Guinea; Australia: Queensland, from near
curling flakes; inner bark orange brown and white; Cape York south to New South Wales.
outer bark grey-brown to dark brown or maroon. TDWG codes: 43 NWG-IJ NWG-PN 50 NSW QLD-
Resin exudate yellowish white, thick and opaque, QU
drying to an amber colour. Crown in mature trees
broadly conical and finally flat-topped, with branch-
Ecology
es only in the top of the tree; branching according
to Massart’s model. Primary first order branches In New Guinea, where this species is widespread, it
of mature trees in pseudo-whorls of 6–10, up to occurs most commonly in montane and high mon-
5 m long, spreading or assurgent, mostly with sec- tane forests above 1000 m altitude, and in cooler
ond to third order branches, caducous. Adventi- and wetter sites than A. hunsteinii. In Australia it
tious foliage branches common. Foliage branchlets is a component of ‘dry vine forest’ and thickets that
assurgent or nearly erect in mature trees, forming occur on the eastern slopes of the Great Dividing
tufts of foliage, of unequal length up to 40 cm, up Range at moderate elevations. In Australia, its range
to 15 mm wide (including leaves), of equal width partially overlaps that of A. bidwillii, and in locali-
but with a tapering apex, more or less flexible, ca- ties where both species occur, they are frequently
ducous. In mature trees leaves of two kinds: adult co-dominant emergents. The two species have dif-
leaves on primary branches and semi-juvenile leaves ferent periods of seed ripening and dispersal. Arau-
on adventitious branches. Adult leaves on ultimate caria cunninghamii is wind dispersed and the seeds
branchlets imbricate or variously spreading but in- ripen in the period overlapping the two monsoons,
curved, subulate, (2–)4–11 × (0.6–)1–3.5(–6) mm, whereas A. bidwillii is animal dispersed and produc-
widest below middle, slightly concave adaxially, es ripe seed between December and April which is
4-faced, keeled on both sides, distally tapering to both the hottest and the wettest period in the Bunya
an incurved, acute or subacute, blunt apex; margins Mountains. In New Guinea, A. cunninghamii and
A. ­hunsteinii also occur together, but usually one Araucaria cunninghamii Aiton ex A. Cunn. var.
species predominates. In New Guinea, common cunninghamii. Type: Australia: Queensland, [“East
tree genera with A. cunninghamii are ­Castanopsis, Coast New Holland 1818–1829”], A. Cunningham
Lithocarpus, Cinnamomum, Calophyllum, and s.n. (lectotype K). Fig. 49
Schizo­meria. In Australia, Argyrodendron is the
most common dominant; numerous other trees and
Description
lianas are found in these forests where the vegeta-
tion is left undisturbed. Araucaria cunninghamii is Juvenile leaves (4–)6–20(–23) mm long, 0.8–2 mm
an emergent rising well above a closed canopy of wide. Microsporophylls usually rhombic, smooth;
­angiosperm trees at around 30 m. apex acute.
199
Conservation Distribution
In Australia, logging by European colonists from the Australia: coastal Queensland, New South Wales
19th century onwards has greatly reduced the area of (Dorrigo Plateau).
occupancy (AOO) of A. cunninghamii, especially TDWG codes: 50 NSW QLD-QU
in New South Wales, where by the beginning of the
20th century only small fragments remained (Baker
Conservation
& Smith, 1910). In Queensland the reduction was
less severe, but here only anecdotal evidence exists IUCN: LC
concerning its extent before European settlement.
Deforestation in the highlands of New Guinea has Araucaria cunninghamii Aiton ex A. Cunn. var.
a much longer history, but, until recently, Papuans papuana Lauterb., Bot. Jahrb. Syst. 50: 51. 1914.
had no incentives to exploit the trees of this species Type: Papua New Guinea: Morobe, Waria River,
on a large scale and in fact often protected individual [“Wasserscheide zwischen Küste und dem Waria-
stands while burning the surrounding (secondary) Inlandtal, Berggrat, 2000 m”], H. Klink 3 (lectotype
grassland. It is therefore difficult to quantify total re- K).
duction of the species within a time frame of ‘three
generations’ (of trees) as required by IUCN Red List
Description
criteria. Due to its large ranges both in Australia and
New Guinea, the species (and its two varieties) is Juvenile leaves 6–20(–22.5) mm long, 1–2.3 mm
considered not threatened at the present time. wide. Microsporophylls usually with a rounded up-
IUCN: See under varieties for ratings. per margin and rugose surface.

Uses Distribution
The timber of Hoop pine is easily worked and is New Guinea (including Ferguson & Japen Islands)
used for joinery, construction, plywood, balus- from the Owen Stanley Range to the Vogelkop
trades, decking, doors, musical instruments, aircraft (Bird’s Head) Peninsula, but especially abundant in
components, etc. There are extensive plantations in the central highlands.
Australia (production capacity 400,000 m3 per year) TDWG codes: 43 NWG-IJ NWG-PN
and also in other parts of the world, e.g. Vanuatu
and India, in particular around Dehra Dun and in
Conservation
the foothills of the Himalaya. This species was intro-
duced to cultivation in 1827 and is commonly grown IUCN: LC
in city parks, e.g. in New Zealand and South Africa,
where some trees have attained large dimensions.

2 varieties are recognized:

 Araucaria heterophylla (Salisb.) Franco, Anais and slightly curved at anthesis, 4–5 cm × 10–15 mm.
Inst. Super. Agron. (Lisboa) 19: 11. 1952. Eutassa Microsporophylls spreading at 45° from the rachis;
heterophylla Salisb., Trans. Linn. Soc. London 8: stalk linear, thin and weak, 3–4 mm long; lamina
316. 1807. Type: Australia: Norfolk Island, P. G. King rhombic, 3–4 × ca. 4 mm; apex acute or apiculate,
s.n. (lectotype G). Fig. 50 upturned. Pollen sacs 10–12, linear, straight, later
curved, 4–5 mm long. Seed cones terminal on 4–10
cm long, very thick foliage branches, usually soli-
Etymology
tary, erect, subglobose to globose, 7.5–10.5 × 8–11.5
The species epithet refers to the very different ju- cm (green weight to 1 kg). Bracts cuneate-flabellate,
venile and adult foliage, in contrast to the homo­ 3.5–4 × 2.5–3 cm including rounded, thin membra-
200 phyllous leaves of A. araucana, which was the only nous wings, distally thickened, ending in a caudate,
other species of the genus known at the time. upward curved, 5–8 mm long tip. Seed scales not
wider than the seed; ligule triangular, 2–3 mm long.
Seeds oblong-cuneate, 2.5–3 × 1.2–1.5 cm, slightly
Vernacular names
flattened, striated longitudinally or smooth, light
Norfolk Island pine, Star pine. brown when ripe.

Description Taxonomic notes

Monoecious trees to 57 (in the past to 70) m tall, to Araucaria heterophylla is the ‘type’ species for Mas-
1.5(–3) m d.b.h.; trunk straight. Bark to 5 cm thick, sart’s model of architecture in trees.
exfoliating in horizontal strips, forming small,
curled flakes; inner bark reddish brown; outer bark
Distribution
light grey. Resin exudate yellow to orange, resem-
bling amber, drying white or brown. Crown in ma- Australia: Norfolk Island, where now largely restrict-
ture trees narrowly conical, branched along greater ed to the Norfolk Island National Park (formerly Mt.
part of trunk; branching according to Massart’s Pitt Reserve). The island’s area is only 34 km2. The
model. Primary first order branches of mature trees species also occurs on Philip Island, a 4 km2 small
in pseudo-whorls of 5–7(–10), 2–3 m long, spreading island 6 km S of Norfolk Island.
horizontally and assurgent at ends or ascending, ca- TDWG codes: 50 NFK-NI
ducous. Adventitious foliage branches occurring in
older trees mostly after damage. Foliage branchlets
Ecology
steeply ascending in mature trees and confined to
distal parts of primary branches (and adventitious Araucaria heterophylla occurs on basalt of the island
branches), 30–40 cm long, 7–12 mm wide (includ- remnant of a Pliocene volcano sitting on the high-
ing leaves), straight or slightly curved, more or less est point of the submerged Norfolk Island Rise, most
terete, flexible. In mature trees leaves of two kinds: abundantly at lower elevation up to ca. 120 m, more
adult leaves on primary branches and semi-juvenile sparsely and remaining smaller up to the highest
leaves on adventitious branches. Adult leaves on ul- point of the island (Mt. Bates, 318 m). The climate
timate branchlets scale-like, ovate or triangular, 4–7 is subtropical with an average rainfall of 1350 mm
× 3–4(–6) mm, 3-faced, showing rhythmic varia- evenly distributed. The original vegetation cover
tion in size along a branch; margins thick and en- has been largely disturbed except near the highest
tire; apex incurved, acute or obtuse, with an abaxial point, hence the present occurence as solitary trees
keel. Stomata on both surfaces, on abaxial surface on coastal headlands or in ‘groves’ with a low un-
proximal only and on either side of the abaxial keel. dergrowth of mostly grasses or of open scrub, with
Semi-juvenile leaves closely adpressed, acicular, very many introduced species. The original vegetation
gently falcately curved for most of their length, with was an evergreen subtropical forest with angiosperm
incurved apex. Pollen cones terminal on ultimate trees and tree ferns 10–20 m high, over which
branchlets, solitary, initially erect but pendulous the araucarias emerged. Common angiosperm
when shedding pollen, obovoid then ovoid-ellipsoid trees are (were) Nestegis apetala, Elaeo­dendron
curti­pendulum, Streblus pendulinus, Lagunaria Araucaria humboldtensis J. T. Buchholz, Bull.
­patersonia, Acronychia simplicifolia, ­Zanthoxylum Mus. Hist. Nat. (Paris), sér. 2, 21: 279. 1949. Type:
pinnatum, Meryta angustifolia, ­Baloghia lucida, and New Caledonia: Grande Terre, Province Sud, Mt.
Dysoxylum patersonianum. Humboldt, J. T. Buchholz 1571 (holotype ILL).

Conservation Etymology
Norfolk Island, although it may have been visited by This species was named after the type locality, Mt.
seafaring Polynesians, was uninhabited when James Humboldt.
Cook found it on his second voyage in October 1774.
At that time the island was covered with a mixed ev- 201
Vernacular names
ergreen rainforest in which A. heterophylla was an
emergent occurring everywhere in greater or lesser No common names have been recorded for this spe-
densities, probably with least density at the highest cies.
elevation points and greatest density on the lower
plateaux nearest the coast. Of this, only scattered
Description
stands of araucarias remain, while the rainforest
has mostly disappeared. Dieback has been observed Monoecious trees 6–20(–30) m tall, to 0.7 m d.b.h.;
recently in some Norfolk Island Pine subpopula- trunk straight. Bark to 3 cm thick, exfoliating in hor-
tions. The natural populations are threatened by in- izontal strips, forming small, curled quadrangular
troduced species such as Lantana camara, Psidium flakes; inner bark brown; outer bark light (yellow-
cattleyanum and Solanum mauritianum. The tiny ish) grey or light green. Resin exudate white, opaque.
population on Philip Island is critically endangered Crown in mature trees columnar, open, branched
(CR) (Greene, 1979, cited in Golte, 1993). along greater part of trunk, usually flat-topped and
IUCN: VU (B1+2c) candelabra shaped; branching according to Mas-
sart’s model. Adventitious foliage branches rare or
absent. Primary first order branches of mature trees
Uses
in pseudo-whorls of 5–7, 1.5–2.5 m long, spread-
Norfolk Island pine is probably the most widely ing horizontally or ascending, caducous on lower
cultivated species of Araucaria. As well as being part of trunk. Foliage branchlets steeply ascending
introduced as a plantation or outdoor ornamen- on primary branches and confined to distal parts,
tal tree in suitable climates, e.g. in Australia, New 5.5–23(–30) cm long, 3–9(–11) mm wide (including
Zealand, Hawaii, Fiji, Micronesia, Pitcairn Island, leaves), more or less the same width throughout
Philippines, South Africa, Côte d’Ivoire, Egypt, except for base, straight or slightly curved, more or
Turkey, Canary Islands, Puerto Rico, Cuba, Florida less terete, flexible. Adult leaves on ultimate branch-
and southern California, it is frequently used world- lets helically arranged obliquely to axis and over-
wide as an indoor ornamental houseplant in offices lapping slightly, scale-like, ovate, 4–6(–7) × 2–4.5
and similar buildings. It is salt tolerant and is con- mm, 4-faced, similar in shape and size throughout
sequently used in landscaping in areas susceptible the length of a branchlet, keeled abaxially; apex in-
to sea spray. However, it is not as wind-resistant as curved, acute. Stomata on both sides, on the abaxial
some other species of Araucaria, especially in ar- side restricted to the proximal part and the extreme
eas that are also susceptible to frosts, to which the apex, on the adaxial side in regular, closely spaced
species is not very resistant. This species has been lines from middle to apex. Pollen cones terminal,
introduced and is now self-sown on Lord Howe Is- subtended by lanceolate, incurved leaves, cylindri-
land beside the lagoon, where it is becoming natu- cal, initially erect, pendulous and slightly curved at
ralised, to the detriment of native species because of anthesis, 3–3.5 cm × 9–10 mm. Microsporophylls im-
its invasive nature and its capability to change the bricate; stalk thin, 3–4 mm; lamina triangular, 3 × 3
soil chemistry, making conditions unsuitable for the mm; apex acuminate to acute. Pollen sacs ca. 6, thin,
germination of many native species. For these rea- straight. Seed cones terminal on short, very thick fo-
sons it is being targeted for eradication there. liage branches, usually solitary, erect, broadly ovoid
to subglobose, 6.5–9 × 4.5–8 cm. Bracts cuneate-
 flabellate, 2–3 × 1.8–2.8 cm including rounded, thin Araucaria hunsteinii K. Schum., in Schumann
membranous wings, distally thickened, ending in & Hollrung, Fl. Kaiser Wilhelms Land: 11, t. 4,
an upward curved, 4–5 mm long tip. Seed scales not f. 8. 1889. Titanodendron hunsteinii (K. Schum.)
wider than the seed; ligule triangular, 1–3 mm long. A. V. Bobrov & Melikyan, Komarovia 4: 61. 2006.
Seeds oblong, 17–22 × 9–10 mm, slightly flattened, Type: Papua New Guinea: Morobe, Butaweng,
striated longitudinally or smooth, light brown when [“oberhalb Bataueng”], C. Hunstein s.n. (holotype
ripe. not located, isotype K).

Araucaria schumanniana Warb., Monsunia 1: 187, t.

Distribution
10, f. A. 1900; Titanodendron schumanniana (Warb.)
202 New Caledonia: Grande Terre, Province Sud, con- A. V. Bobrov & Melikyan, Komarovia 4: 61. 2006.
centrated in the large southern massifs (Massif du Araucaria klinkii Lauterb., Bot. Jahrb. Syst. 50: 48,
Humboldt, Montagne des Sources, Mt. Mou). f. 1. 1914; Araucaria hunsteinii K. Schum. var. klinkii
TDWG codes: 60 NWC (Lauterb.) Silba, Phytologia 68: 26. 1990; Titano­
dendron klinkii (Lauterb.) A. V. Bobrov & Melikyan,
Komarovia 4: 61. 2006.
Ecology
This species occurs in montane dense humid forest
Etymology
and especially in montane tall maquis (2–8 m), on
ultramafic soils derived from serpentine or peri­ Named after Carl Hunstein (ca. 1843–1888), German
dotite. It reaches to the ridges and summits of the collector in New Guinea in 1878–1888, after whom
highest mountains of the ‘Grand Massif du Sud’ and the Hunstein Range in East Sepik in Papua New
is especially abundant on E and SE slopes and in gul- Guinea is also named.
lies. Altitudinal range: 750–1600 m a.s.l. It is some-
times associated with A. montana or A. laubenfelsii
Vernacular names
and often with other conifers such as Callitris neo-
caledonica, Falcatifolium taxoides, Prumnopitys fer- Klinki pine (also trade name for timber). There are
ruginoides, and Podocarpus spp., and numerous many local names in New Guinea for this tree.
angiosperm trees and shrubs, e.g. Metrosideros sp.
(Myrtaceae), Araliaceae, Dilleniaceae and Lauraceae.
Description
Tree ferns (Dicksonia deplanchei) and small palms
are also often present, while ferns and sedges form a Monoecious trees to 90 m tall, to 2 m d.b.h. or more;
herbaceous layer. Lichens and mosses are very abun- trunk straight, very gradually tapering. Bark thick,
dant, especially at the higher altitudes above 1200 m. rough and scaly, exfoliating in large plates and scales;
inner bark reddish; outer bark dark brown to black.
Resin exudate clear, changing to golden-coloured on
Conservation
contact with air and weathering white to grey. Crown
This species mainly occurs on remote mountain in mature trees very open, with several branches
massifs that are (partly) in nature reserves and at retained in lower part of tree, flat-topped; branch-
present not threatened by mining activities although ing according to Rauh’s model. Adventitious foliage
the geology is similar to the ultramafic rock types branches common in older trees. Primary first order
that are actively mined. The main threats are fre- branches of mature trees in pseudo-whorls of 6–8,
quent fires, some of which are very destructive and up to 6 m long, spreading horizontally, bearing foli-
difficult to fight. age branches near distal end, slowly caducous. Foli-
IUCN: EN [B1ab(iii)+2ab(iii)] age branchlets spreading in all directions or pendent
on ends of primary branches in mature trees, form-
ing large tufts of up to 50 cm long foliage branchlets
Uses
of very unequal width (including leaves) from base
The wood is not exploited and there are no other re- to apex. In mature trees leaves of two kinds: adult
corded uses. leaves on primary branches and semi-juvenile leaves
on adventitious branches. Adult leaves of ultimate
branchlets imbricate at base but spreading radially are merely the adult stage of A. hunsteinii. With the
at a wide angle, often in 5 distinct rows, bifacially availability of more specimens we can be sure now
flattened, triangular to lanceolate, (2–)5–10(–15) cm that Lauterbach based his species on mature foliage
× (8–)12–20(–25) mm, increasing in size from base of A. hunsteinii and that therefore A. klinkii is a taxo-
to apex of branchlet; margins tapering to a sharply nomic synonym of A. hunsteinii. Forms with glau-
pungent apex. Stomata in numerous intermittent cous blue-green leaves and male and female cones
rows covering both leaf surfaces from base to apex. with white exudate or powder on their surface have
Leaves on adventitious branches variable in length been called A. klinkii Lauterb.; however, De Lauben-
but narrower than adult leaves. Pollen cones axillary, fels (1988) has pointed out that the young leaves are
sessile or pedunculate, in clusters of 2–6, narrowly glaucous, and in some forms the adult leaves and
cylindrical, initially erect, elongated, pendulous and female cones are grey-blue due to a white exudate. 203
curved at anthesis, 10–22 cm × 11–25 mm. Micro-
sporophylls spreading at ca. 70° from a thin, frag-
Distribution
ile rachis; stalk thin and weak, linear, 4–5 mm long;
lamina curved, the exposed part ligulate, 5–8 × 2.5–3 Papua New Guinea. In the central mountain range
mm; abaxial surface rugose, lateral margins erose- from an isolated population on the Wamira River in
denticulate; apex acuminate. Pollen sacs 8–10, linear, the east through the Owen Stanley Range and the
straight, ca. 6 mm long. Seed cones axillary (appear- Bismarck Range, with other isolated stands near
ing subterminal) on a short, stout, leafy peduncle, Sattelburg in the Huon Peninsula and on the Tagari
usually solitary, erect, broadly obovoid to cylindri- River in the Central Highlands.
cal, 15–25 × 12–16 cm. Bracts obtrullate to cuneate, TDWG codes: 43 NWG-PN
5–6 × 7–9 cm including wings, lateral sections thin
and membranous, distal part thickened to a rhom-
Ecology
bic, sharply keeled apophysis, ending in a 8–15 mm
long, caducous tip. Seed scales not wider but longer This species forms ‘groves’ or stands, or occurs scat-
than the seed; ligule small, 2–3 × 5 mm. Seeds nar- tered in tropical montane monsoon forests on moist
rowly almond-shaped, ca. 30 × 8 mm, distinctly flat- sites usually in inter-montane valleys. Its altitudi-
tened, smooth. nal range is (550–)750–1700(–2100) m a.s.l. Ma-
ture trees are canopy emergents above angiosperm
trees, effectively forming an upper canopy layer.
Taxonomic notes
This species occurs in two types of forest, a drier
The two species of Araucaria that are indigenous in and a wetter one; in the drier type the canopy of an-
New Guinea, A. hunsteinii and A. cunninghamii, are giosperms reaches only 15–25 m of average height,
distinct and classified in two sections: Intermedia with A. hunsteinii attaining twice that height. As-
White and Eutacta Endl. When Schumann first de- sociated common evergreen tree species in this
scribed A. hunsteinii, he does not seem to have had forest type are Aleurites moluccana, Celtis sp., Her-
a mature seed cone but only an immature cone and itiera sp., Macaranga sp., and Pouteria luzonensis;
some separate bract-scale complexes. Warburg de- deciduous species are Garuga floribunda, Protium
scribed A. schumanniana as a new species distinct macgregorii, Sterculia sp., and Terminalia sp. In high
from A. hunsteinii, but the differences he gave in the rainfall localities both angiosperms and A. hunstei-
description and in the Figures A and B of Plate 10 do nii become much taller, with the conifer towering
not hold when more specimens with mature, full- to 60–90 m. Common associated canopy tree spe-
grown male and female cones are examined. Lau- cies are ­Acmena sp., Elmerillia papuana, Flindersia
terbach described yet another species of Araucaria amboinensis, F. pimenteliana, Pometia pinnata, and
from New Guinea, A. klinkii, a name that apparently Xantho­phyllum papuanum. Cerbera floribunda,
became popularized as the vernacular name for the Cryptocarya sp., Dysoxylum sp., Gnetum gnemon,
species now known as A. hunsteinii. Lauterbach Litsea sp., and Myristica sp. are common in the sub-
himself expressed some doubt about the distinc- canopy tree layer. Soils are neutral to acidic with a
tiveness of his new species from A. hunsteinii and high clay content. Precipitation ranges from 800
considered it possible that the leaves of A. klinkii mm to more than 4000 mm per year.
 lower on the trunk. Primary first order branches of
Conservation
mature trees in pseudo-whorls of 4–5, 1–2.5 m long,
This species was graded LR(nt) by Farjon & spreading nearly horizontally to assurgent, cadu-
Page (1999), which was still the rating (NT) in cous. Foliage branchlets assurgent or nearly erect
the 2008 IUCN Red List of Threatened Species™ on primary branches in mature trees and confined
(www.iucnredlist.org) although the destruction of to distal parts, of more or less equal length (35–45
so many trees in recent forest fires may mean that cm) on either side of the axis branch and shortening
the species needs to be reassessed. The recalcitrant towards the end, (12–)20–33 mm wide (including
seeds cannot be stored for long periods making leaves). On mature trees leaves of two kinds: adult
propagation from seed difficult. leaves and semi-juvenile leaves on adventitious
204 IUCN: NT branches. Adult leaves on ultimate branchlets nearly
triangular, 8–20(–24) × 4–12 mm, at least twice as
long as wide, similar in size along length of branch-
Uses
let, strongly convex, usually with 5 longitudinal ridg-
Araucaria hunsteinii has been grown as a plantation es alternating with grooves or sometimes smooth;
crop in New Guinea since 1948. The wood is used margins incurved; apex acute or obtuse. Stomata
in the local sawmilling and plywood industries and abaxially in bands located in the grooves between
for making aircraft frames. Fires destroyed much the ridges; adaxially in numerous continuous rows
of the Bulolo plantation in 1997. Klinki pine plan- from base to apex. Semi-juvenile leaves irregularly
tations have also been established on tin tailings in spreading at 40–90°, acicular to lanceolate, 13–25 ×
Malaysia and more recently in Costa Rica. The lat- 4–8 mm, tapering to an incurved, acute apex. Pollen
ter plantations were set up as part of a controversial cones terminal, solitary, initially erect but becoming
programme, KLINKIFIX, which aims to mitigate pendulous and curved at anthesis, 8–11(–15) × 2–3
greenhouse gas emissions. cm. Microsporophylls imbricate, spreading at 45°
from a slender rachis; stalk thin and weak, 3–5 mm
long; lamina nearly triangular, 4–6 × 3–5 mm, thick
Araucaria laubenfelsii Corbasson, Adansonia, sér. and firm; lateral margins entire; apex obtuse; abaxial
2, 8: 467. 1969. Type: New Caledonia: Grande Terre, surface smooth. Pollen sacs 10–12, thin, straight, 5–6
Province Sud, on ridge between Mts. Dzumac and mm long. Seed cones terminal on short, stout foliage
Mt. Ouin, H. S. MacKee 16441 (holotype P). Fig. 51 branches, usually solitary, sometimes 2–3 together,
erect, broadly ovoid to subglobose, 10–13 × 7–9
cm. Bracts flabellate, 2.2–2.5 × 1.5–2 cm including
Etymology
rounded, thin membranous wings, distally thick-
This species was named after David J. de Lauben- ened and transversely keeled, ending in a rostrate,
fels, specialist on gymnosperms especially the Podo­ upturned, 10–15 mm long, acute tip. Seed scales not
carpaceae and the Araucariaceae. wider than the seed; ligule fragile, 1.5–2 mm long.
Seeds cuneate, 14–16 × 6–8 mm, smooth, yellowish
brown when ripe.
Vernacular names
ouaoué (name recorded on H. S. MacKee 30370, P).
Distribution
New Caledonia: Grande Terre. In Province Nord
Description
only known from Île Art and Mt. Kaala; more abun-
Monoecious trees to 30 m tall, to 1 m d.b.h.; trunk dant in Province Sud.
straight. Bark to 4 cm thick, exfoliating in horizontal TDWG codes: 60 NWC
strips; inner bark reddish to pink; outer bark brown
or grey. Resin exudate clear or yellow, drying white
Ecology
or grey. Crown in mature trees columnar or broadly
conical, open, branched along the greater part of the Araucaria laubenfelsii is an emergent from om-
trunk; branching according to Rauh’s model. Pri- brophilous lower to middle-montane forest, and oc-
mary branches followed by adventitious branching curs also in ‘maquis minier’; on ultramafic substrates
derived from serpentine or peridotite, or on lateritic
Description
soil at lower elevations. Its altitudinal range is (26–)
250–1160 m a.s.l. On slopes and summit ridges it is Monoecious trees to 40 m tall, to 0.6 m d.b.h.; trunk
often associated with A. montana and sometimes straight or curved. Bark to 2 cm thick, exfoliating
with A. humboldtensis; Myrtaceae, Casuarinaceae, in horizontal strips; inner bark dark red; outer bark
small palms and the conifers Falcatifolium taxoides grey. Resin exudate from bark red, from foliage
and Prumnopitys ferruginoides are common in a sec- branchlets whitish translucent and semi-opaque.
ond tree layer or in the understorey. Regeneration Crown in mature trees variable, often short, densely
is most successful in open and low vegetation and branched, with a conical or rounded top; branching
diminishes when the vegetation closes to become according to Massart’s model. Primary branches fol-
dense forest, in which the surviving araucarias be- lowed by some adventitious branching. Primary first 205
come large and tall emergents. It appears therefore order branches of mature trees in pseudo-whorls of
that this species is dependent on episodic fires for its 5–7 at regular intervals, 2–3 m long, spreading or
perpetuation in the vegetation. assurgent, mostly persistent. Foliage branchlets as-
surgent on primary branches in mature trees and
confined to distal parts, giving a candelabra appear-
Conservation
ance, 7–30 cm long and at thickest parts 9–13(–16)
The threats are those common to most of the New mm wide (including leaves), narrowed at base and
Caledonian species: fires, land management of non- at intervals rhythmically along a branchlet, there
agricultural areas, and clear-cutting. Most popula- only 6–8 mm wide (including leaves). Adult leaves
tions occur on serpentine and some, especially in on ultimate branchlets ovate-lanceolate, ovate or
the Province Nord, are subject to nickel mining. The broadly ovate, 4–16 × 2.5–6.5(–8) mm, slightly con-
species has probably now disappeared from one of vex, abruptly narrowed at base, straight or incurved,
such sites, Mt. Kaala (pers. observation, Nov 2005). loosely imbricate and slightly divergent, strongly
A reassessment of its status seems necessary. keeled abaxially; apex rounded or subacute, strongly
IUCN: NT incurved. Stomata abaxially in intermittent rows
from base to apex or (in smaller leaves) only near
base; adaxially in numerous rows. Semi-juvenile
Uses
leaves on adventitious branches 6–10 × 0.7–2.5(–3.2)
The wood of this species is not exploited and no mm. Pollen cones terminal, solitary, initially erect
other uses are recorded. but pendulous and curved at anthesis, (10–)12–15
(–17) cm × 20–28 mm. Microsporophylls imbricate,
spreading at 45° from a stout rachis; stalk thin and
Araucaria luxurians (Brongn. & Gris) de Laub., weak, 4–5 mm long; lamina ovate-oblong, 8–10 ×
Fl. Nouv. Calédonie Dépend. 4: 92. 1972. Araucaria 5–6 mm, lateral margins denticulate, scarious; apex
cookii R. Br. ex Lindl. var. luxurians Brongn. & Gris, acute when mature. Pollen sacs 12–15, linear, thin,
Ann. Sci. Nat. Bot., sér. 5, 13: 354. 1871; Araucaria straight, ca. 8 mm long, directed towards the rachis.
columnaris (G. Forst.) Hook. f. luxurians (Brongn. Seed cones terminal on short, stout foliage branches,
& Gris) E. H. Wilson, J. Arnold Arbor. 7: 84. 1926. usually solitary, erect, subglobose, 10–12 × 8–10 cm.
Type: New Caledonia: Grande Terre, Province Sud, Bracts flabellate, 3–3.5 × 3–3.5 cm including round-
Mt. Canala, B. Balansa 2510 (lectotype P). ed, thin membranous wings, distally thickened but
thin-edged, ending in a caudate, straight or upward
curved, 8–10 mm long tip. Seed scales not wider
Etymology
than the seed; ligule triangular, 2–3 mm long. Seeds
The species epithet (Latin luxurians = ‘luxuriant’) obovoid, 18–25 × 6–10 mm, more or less flattened,
refers to the branching habit. smooth, light brown when ripe.

Vernacular names Taxonomic notes

sapin de Noël (New Caledonia) Araucaria luxurians was first described by Brongni-
art & Gris (1871) as a new variety of Araucaria cookii
 (= A. columnaris) and it was said to occur along the Some localities, like Mt. Kaala and Col de Mô, were
coast mixed with that species. The authors described revisited in November 2005 by me, and no trees of
the variety as having larger, roundly ovate leaves 8–9 A. luxurians could be found there at that time. The
mm long and having larger, often curved pollen most recent assessment estimated fewer than 2500
cones being 12 cm long with larger pollen sacs. Ac- mature trees.
cording to De Laubenfels (1972), who first raised it IUCN: EN [B1ab(iii);C2a(i)]
to the rank of species, the two taxa are ecologically
distinct, with A. columnaris growing on limestone
Uses
(coral) and A. luxurians on serpentine. However, De
Laubenfels’s morphological justification is minimal. The wood is of high quality and has been locally ex-
206 On the same branchlet, leaves mostly vary with each ploited. It is a beautiful tree, especially in younger
burst of growth. Adult leaves, being either smaller stages, and is used as an ornamental on ultrabasic
or larger than those of A. columnaris, can easily be soils in New Caledonia.
found on a single tree of A. luxurians. The pollen
cones of A. luxurians are indeed about twice, some-
times nearly three times as large as those of A. co- Araucaria montana Brongn. & Gris, Nouv. Arch.
lumnaris; its constituent organs are therefore also Mus. Hist. Nat. Paris 4: 215, t. 14. 1868. Type:
larger. The habit of the smaller species A. luxurians New Caledonia: Grande Terre, [“Sommet Mi”],
is less columnar than in A. columnaris and reitera- B. Balansa 2512a (lectotype P).
tion of its first order branches, which are persistent,
is rare. Araucaria luxurians is here accepted as a dis-
Etymology
tinct species.
The species epithet means “of the mountains”.
Distribution
Vernacular names
New Caledonia: Grande Terre and Îles Bélep.
TDWG codes: 60 NWC None are recorded.

Ecology Description
Araucaria luxurians is forming escarpment forest Monoecious trees to 40 m tall, to 1 m d.b.h.; trunk
on serpentine cliffs next to the sea or in the inte- straight. Bark to 3 cm thick, exfoliating in horizontal
rior, with an altitudinal range of 1–1000 m a.s.l. It strips; inner bark red or dark red; outer bark pale
grows in rocky ravines, or on coastal mountains and grey, sometimes almost white. Resin exudate white.
there usually in ‘maquis minier’; on eroded, ultra- Crown in mature trees broadly domed, very open,
mafic substrates of serpentine or peridotite origin. branched in the upper part of trunk, flat-topped in
De Laubenfels (1972) gives the substrate for this older trees; branching according to Rauh’s mod-
species as on serpentine; in some lowland coastal el. Primary first order branches of mature trees in
areas where it may be more or less sympatric with pseudo-whorls of 4–5, 1–3 m long, spreading near-
A. columnaris; the latter species is mostly restrict- ly horizontally to assurgent, caducous. Adventi-
ed to limestone derived from coral reefs, whereon tious foliage branches similar to primary branches
A. luxurians is not known to occur. The Bélep isles but shorter. Foliage branchlets assurgent or nearly
are of peridotite (Golte, 1993). erect in mature trees and confined to distal parts of
primary branches (and on adventitious branches),
often curved, 15–45 cm long, (12–)15–25 mm wide
Conservation
(including leaves), more or less flexible. On mature
According to the 2008 Red List of Threatened Taxa™ trees leaves of two kinds: adult leaves on primary
(www.iucnredlist.com), there are fewer than five branches and semi-juvenile leaves on adventitious
populations still extant, none of which is protected. branches. Adult leaves on ultimate branchlets broad-
Some are seriously threatened by fire and erosion, ly ovate, 8–15 × 6–11 mm, similar in size throughout
while settlement and mining are other concerns. the length of the branchlet, strongly incurved near
the obtuse apex, abaxially with longitudinal grooves
Uses
or striate, more or less obscurely keeled. Stomata
on the abaxial surface restricted to the lower half of The wood is not exploited. It has been planted in ec-
the leaf, more numerous on the adaxial face. Semi- ological restoration projects in New Caledonia. No
juvenile leaves 5–12 × 4–7 mm, arcuate, with an other uses are recorded.
abaxial keel; apex incurved. Pollen cones terminal,
solitary, erect and remaining straight, cylindrical,
5–16.5 cm × 15–30 mm. Microsporophylls spread- Araucaria muelleri (Carrière) Brongn. & Gris,
ing at 45° from a stout rachis; stalk thin and weak, Nouv. Arch. Mus. Hist. Nat. Paris 4: 219, t. 15 & 16.
2–3 mm long; lamina ovate-triangular, with a thick, 1868. Eutacta muelleri Carrière, Rev. Hort. 37: 392.
curved base, 5–7 × 4–5 mm; lateral margins minute- 1866. Type: Illustration in Rev. Hort. 37, pl. inserted 207
ly denticulate or papillate; apex acute; abaxial sur- betw. pp. 392–393, f. 3. 1866 (lectotype). Fig. 52
face smooth. Pollen sacs 10–12, linear, straight, 5–6
mm long. Seed cones terminal on short, stout foliage
Etymology
branches, usually solitary, erect, broadly ovoid to
subglobose, 8–11 × 7–9 cm. Bracts flabellate, 2.5–3.5 Named after Ferdinand Jacob Heinrich von Mueller
× 1.8–2.5 cm including rounded, thin membranous (1825–1896), a famous German botanist who studied
wings, distally thickened and transversely keeled, the flora of Australia.
ending in a caudate, upturned, 10 mm long, acute
tip. Seed scales not or slightly wider than the seed;
Vernacular names
ligule small, triangular, fragile, 1.5–2 mm long. Seeds
cuneate, 20–25 × 7–8(–10) mm, smooth, yellowish No common names have been recorded for this spe-
brown when ripe. cies.

Distribution Description
New Caledonia: Grande Terre. Monoecious trees to 15(–20) m tall, to 0.6 m d.b.h.;
TDWG codes: 60 NWC trunk straight. Bark to 3 cm thick, exfoliating in hor-
izontal strips; inner bark dark red; outer bark light
grey or reddish grey. Resin exudate translucent to
Ecology
opaque, drying white. Crown in mature trees very
Araucaria montana occurs in dense and low humid open, candelabra shaped, branches along greater
forest and in tall ‘maquis minier’ (‘maquis buisso- part of trunk but few in number; branching accord-
nant’) on mountain ridges, in ravines and on elevated ing to Rauh’s model. Primary first order branches of
plateaux, often on ultramafic peridotite, serpentine mature trees 1.5–3 m long, bearing foliage branches
or brown laterite, more rarely on acidic micaschists. towards the ends, several branches lowest on the
Altitudinal range: (200–) 500–1400 m a.s.l., but usu- trunk caducous. Adventitious foliage branches rare
ally in the range 800–1200 m. It is sometimes a co- or absent. Foliage branchlets steeply ascending to
dominant in Araucaria-Nothofagus forest. erect on primary branches in mature trees and con-
fined to distal parts, to 50 cm long, 3–5 cm wide
(including leaves), narrowest proximally and broad-
Conservation
est distally, straight or slightly curved, more or less
This is a widely distributed species and one of the terete, rigid. Adult leaves on ultimate branchlets lan-
few that occurs not exclusively on ultramafic sub- ceolate to ovate, 20–32 × 9.5–18.5 mm, flat or weakly
strates. Populations are often quite extensive, yet it is navicular, widest just above base, tapering to an ob-
affected by nickel mining in several of its locations tuse apex; abaxial surface coriaceous, smooth, typi-
and expansion of mining threatens a continuing re- cally matt but rarely very lustrous (these forms also
duction of numbers of mature trees and ultimately ± lacking abaxial stomata), with strong slightly ex-
populations. centric longitudinal abaxial keel from base to apex;
IUCN: VU [A3c; B1ab(iii)] margins inrolled. Stomata on both faces of leaves in
 numerous intermittent, closely spaced rows from Regeneration has been observed in the field, but
base to apex, in some trees with lustrous abaxial leaf seems to be slow and could not stand up against
surfaces nearly absent on that face. Pollen cones ter- increased incidence of fires, which are particularly
minal on the ultimate foliage branches, erect when frequent in the far south. This is for this species the
shedding pollen, straight, 20–25 × 3–4 cm. Micro- main threat, due to its predominant habitat ‘maquis
sporophylls attached to a stout, 10–12 mm thick minier’. Many populations are considered to be se-
rachis, imbricate, the laminate part spreading but verely fragmented as a result of fires.
with an incurved apex; stalk linear, 4–5 mm long; IUCN: EN [B1ab(iii)+2ab(iii)]
lamina broadly ovate, 6–8 × 5–6 mm, thick near
the base; margins entire or erose-denticulate; apex
Uses
208 mucronate. Pollen sacs 15–20, linear, ca. 10 mm long,
strongly contorted after dehiscence. Seed cones ter- This species has been used in local reforesting and
minal on short, very thick branches, usually solitary, ecological restoration projects, e.g. at Chûtes de la
erect, broadly ovoid, 11–15 × 9–11 cm. Bracts flabel- Madeleine.
late, 2–2.5 × 1.8–2.2 cm including rounded, thin
membranous wings, distally thickened, ending in
a rostrate, upward curved, 10–15 mm long tip. Seed Araucaria nemorosa de Laub., Trav. Lab. Forest.
scales not wider than the seed; ligule triangular, 2–3 Toulouse T. 1 (8, 5): 1. 1969. Type: New Caledonia:
mm long. Seeds oblong, 17–22 × 9–10 mm, slightly Grande Terre, Province Sud, Port Boisé, western
flattened, light brown when ripe. shore, H. S. MacKee 20218 (holotype P).

Distribution Etymology
New Caledonia: Grande Terre, Province Sud. The species epithet is from Latin nemorosa, pertain-
TDWG codes: 60 NWC ing to woods and groves.

Ecology Vernacular names

Araucaria muelleri occurs both in maquis and open No common names have been recorded for this spe-
thickets above a shrub layer and in dense humid cies.
forest with Nothofagus, but most commonly in
‘maquis buissonant’ and ‘maquis minier’, on serpen-
Description
tine ridges, peridotite, gabbros and iron-rich rocks
(‘ironstone’ or ‘cuirasse de fer’) or brown laterite. Monoecious trees to 20 m tall, to 0.7 m d.b.h.; trunk
Its altitudinal range is 150–1000(–1250) m a.s.l. In straight. Bark to 2 cm thick, exfoliating in horizon-
lowlands this species is commonly associated with tal short strips and flakes; inner bark dark red; outer
Dacrydium araucarioides (Podocarpaceae) and bark grey. Resin exudate translucent, drying white.
Gymnostoma chamaecyparis (Casuarinaceae), both Crown in mature trees variable, columnar or domed
of which have nearly the same candelabra shaped or umbrella-shaped, if columnar then flat-topped;
habit which is also assumed by A. muelleri on a larg- branched mainly in the upper half of the trunk;
er scale. Populations are invariably small and consist branching according to Massart’s model. Primary
of scattered individuals, often at considerable dis- first order branches of mature trees in pseudo-whorls
tance from each other. of 5–7, 1–2.5 m long, spreading, assurgent, mostly
persistent. Adventitious foliage branches on the low-
er part of the trunk, shorter than primary branches.
Conservation
Foliage branchlets assurgent to erect on primary
This species has been found in more localities since branches and confined to distal parts, in two pecti-
De Laubenfels (1972) published his map, but it can- nate rows, 10–22 cm long, rhythmically narrowed at
not be concluded from this that the species has in- intervals along a branchlet, thicker parts 6–12 mm
creased. Populations are almost always very small, wide, thinner parts 2.5–6(–7) mm wide (including
often down to 1–10 trees in a particular location. leaves). Adult leaves on ultimate branchlets variable,
somewhat spreading, subulate-acicular to narrowly
Conservation
lanceolate, 4–10 × 1.3–3 mm, keeled abaxially; apex
incurved, obtuse or acute; leaves subtending pollen This species is among the most seriously threatened
cones distinct, spreading, more or less linear, 10–15 of the 19 species of Araucaria. Two distinct localities
× 1.5–2.5 mm, widest at base. Stomata abaxially in are known to exist with certainty, in the main local-
6–7 intermittent rows each side of keel; adaxial sto- ity (Port Boisé) several subpopulations exist, while
mata numerous. Semi-juvenile leaves on adventi- at Cap Reine Charlotte there is only one very small
tious branchlets 4-angled, acicular, 3.5–9 × 0.5–1.8 population. The potential for fire is very high in the
mm. Pollen cones terminal, solitary, initially erect area due to the type of vegetation and increased
but pendulous and curved near base at anthesis, activities such as road building and mining. None
narrowly cylindrical, 5–12 cm × 10–20 mm. Micro­ of the (sub)populations of this species are in a pro- 209
sporophylls with laminar parts spreading at nearly tected area and they occur just outside the narrow
90° from a stout rachis at anthesis; stalk thin and belt of A. columnaris which benefits from traditional
weak, 5–7 mm long; lamina ovate, 5–8 × 4–5 mm; protection by the Kanaks. The land is mostly pri-
lateral margins denticulate; apex acute or acumi- vately owned by a few Kanak residents of Port Boisé.
nate. Pollen sacs linear, ca. 12, directed towards the Expansion of human settlement and possibly tour-
rachis, ca. 8 mm long. Seed cones terminal on short, ism are other threats to the population.
stout foliage branches, usually solitary, erect, ovoid IUCN: CR [B1ab(iii)+2ab(iii)]
to subglobose, 8–12 × 6–10 cm. Bracts flabellate,
2.6–3 × 2.3–2.6 cm including rounded, thin mem-
Uses
branous wings, distally thickened but thin-edged,
ending in a rostrate, upward curved, 11–15(–20) mm None are recorded.
long tip. Seed scales not wider than the seed; ligule
triangular, 2–3 mm long. Seeds obovoid, 20–22 ×
7–9 mm, smooth, brown when ripe. Araucaria rulei F. Muell., Rep. Burdekin Exped.:
18–19. 1860. Type: New Caledonia: Grande Terre,
[locality not given], W. H. Duncan s.n. (lectotype
Distribution
K).
New Caledonia: Grande Terre, Province Sud, near
Port Boisé and nearby at Cap Reine Charlotte.
Etymology
TDWG codes: 60 NWC
Named after John Rule, who was a nurseryman in
Melbourne, Victoria, in the mid-19th century.
Ecology
The main population of this species occurs on a
Vernacular names
level plateau just inland from Port Boisé at around
20–40 m a.s.l. which consists of impoverished in- No common names have been recorded for this spe-
durated ferritic soils overlying peridotites. The alti- cies.
tudinal range of A. nemorosa is (1–)20–50(–100) m.
The trees stand in small groups or wide apart among
Description
low (to 10 m) and open to dense evergreen forest
or scrub interspersed with nearly black, stony areas Monoecious trees to 20(–25) m tall, to 0.6 m d.b.h.;
(cuirasse) almost devoid of vegetation. Common trunk straight, sometimes curved. Bark to 2 cm
angiosperm species here are Baeckea ericoides and thick, exfoliating in horizontal strips; inner bark
Melaleuca quinquenervia and abundant Gymnos- dull brown or dark red-brown; outer bark light grey.
toma chamaecyparis. Where the plateau falls off to Resin exudate copious, white. Crown in mature
the bay A. nemorosa is replaced by A. columnaris, trees very open; branches along greater part of trunk
which forms dense stands lining the steep coastal but few in number, branching according to Rauh’s
strip and narrow beaches as well as the headlands. model. Primary first order branches of mature trees
Annual precipitation at Port Boisé is approximately 1.5–3 m long, middle and upper ones assurgent but
2500–3000 mm. lower ones strongly declinate and turned upwards
 only at the end; most branches finally caducous. larger populations extend over several km2, such
Adventitious foliage branches rare, usually on other as the one SE of Baie de Poro (Néjeré Dumwà); in
branches. Adult leaves on ultimate branchlets broad- other places only a few trees are present. They are
ly lanceolate, (8–)13–25 × 5–14 mm, flat, widest and almost always scattered and do not form closed for-
curved just above base, tapering to a curved, acute est stands.
apex, weakly keeled abaxially, smooth or sometimes
striate, lustrous green. Abaxial stomata only in the
Conservation
most proximal part of the leaf, ± obscured by over-
lapping leaves; adaxial stomata in ca. 35 rows from This species is fairly widespread, but unfortunately
base to apex. Pollen cones terminal, erect when often occurs on sites that are subject to extensive
210 shedding pollen but horizontal when old, straight nickel mining operations, which disturb all vegeta-
or curved, 8–25 cm × 20–35 mm. Microsporophylls tion. One of the largest populations SE of Baie de
attached to a stout, 10 mm thick rachis, imbricate, Poro (Néjeré Dumwà), with many hundreds of trees,
the laminate part spreading at 45° with a free apex; is located in such a mining area which has destroyed
stalk linear, 3–7 mm long; lamina broadly lanceolate many trees. The trees, although regenerating, grow
to caudate, 6–9 × 3–5 mm; margins entire or weakly slowly, while the destruction of their habitat is com-
denticulate; abaxial surface smooth; apex acute or plete when it has been stripped of all topsoil and
pungent. Pollen sacs 10–16, linear, 5–8 mm long. much of the overburden of laterite to reach the min-
Seed cones terminal on short, very thick branches, eral-richer bedrock underneath. Fires are also a haz-
usually solitary, erect, broadly ovoid, 8–12 × 5.5–8 ard to this species as their incidence often increases
cm. Bracts flabellate-cyathiform, 2–2.3 × 1.8–2 cm with access provided by mining roads. Few popula-
including rounded, thin membranous wings, dis- tions are at present within protected areas.
tally thickened, ending in a caudate, upward curved, IUCN: EN [A2acd, B2ab(ii,iii)]
15–20 mm long tip. Seed scales not wider than the
seed; ligule triangular, 1–2 mm long. Seeds oblong,
Uses
slightly narrowing towards base of scale, 16–20 ×
9 mm, slightly flattened, striated longitudinally or This species is used in local ecological restoration
smooth, brown when ripe. projects, in which good results have been obtained.
The wood is not exploited. Reports that it was in
cultivation in Britain (Dallimore & Jackson, 1966),
Distribution
including cultivars, cannot be substantiated for lack
New Caledonia: Grande Terre, scattered throughout of herbarium specimens and, in view of the limited
the island but more common in the southern half. knowledge the authors had of New Caledonia and
TDWG codes: 60 NWC its conifers, are likely to have pertained to another
species.
Ecology
This species occurs most often in ‘maquis minier’ Araucaria schmidii de Laub., Trav. Lab. Forest.
on ultramafic soils derived from serpentine, peri- Toulouse T. 1 (8, 5): 1. 1969. Type: New Caledonia:
dotite or ironstone (‘cuirasse de fer’), which may Grande Terre, Province Nord, Mt. Panié, M. Schmid
have weathered to brown laterite, and are very low 858 (holotype P).
in phosphorus and often contain nickel. Its altitu-
dinal range is (150–)400–800(–1150) m a.s.l. The
Etymology
landscape is often described as (degraded) maquis
with forest remnants in ravines, but Araucaria rulei This species was named after Maurice Schmid, who
does not occur in dense rain forest. It can be associ- collected the holotype.
ated with Agathis ovata, another larger conifer that
dominates areas with shrubs rather than trees. A few
Vernacular names
collections were made in “forest”, which would have
been low evergreen forest that can be found in plac- No common names have been recorded for this spe-
es with slightly better soil conditions. Some of the cies.
 TDWG codes: 60 NWC
Description
Monoecious trees to 30 m tall, to 0.8 m d.b.h.; trunk
Ecology
straight or curved, sometimes forked from near
base and occasionally with three or as many as six Araucaria schmidii grows in dense humid montane
trunks. Bark to 2.5 cm thick, exfoliating in hori- cloud forest on steep slopes and exposed ridges at
zontal strips; inner bark reddish; outer bark grey. the edge of the summit plateau of Mt. Panié, on mi-
Resin exudate not observed. Crown in mature trees caschist substrate and often directly from rock crev-
columnar, branched along greater part of trunk, ices. Its altitudinal range is 1400–1630 m a.s.l. This is
flat-topped; branching according to Massart’s mod- the only New Caledonian Araucaria that grows ex-
el. Primary first order branches of mature trees in clusively on non-ultrabasic substrates. At its lowest 211
pseudo-whorls of 5–7, 1–2 m long, spreading or as- limit on the mountain this species is still accompa-
cending, slowly caducous beginning with branches nied by Agathis montana, whereas near the summit
lowest on the trunk. Adventitious foliage branches it is the only tall tree, which is emergent above a
shorter than primary branches. Foliage branchlets dense undergrowth of evergreen angiosperm shrubs
strongly ascending or assurgent on distal parts of and small trees, as well as tree ferns (Cyathea vieil-
primary branches, of unequal length (7.5–25 cm), lardii) and small palms. Mt. Panié’s eastern slopes
in two pectinate rows, shortening towards the end, receive the highest rainfall on the island, estimated
6–10 mm wide (including leaves), flexible; adventi- to exceed 8000 mm per year (Schmid, 1989; cited in
tious branchlets irregularly inserted and arranged. Golte, 1993).
On mature trees leaves of two kinds: adult leaves
and semi-juvenile leaves on adventitious branch-
Conservation
es. Adult leaves on ultimate branchlets subulate-­
lanceolate, 5–9 × 1.2–2 mm, convex, 4-faced, keeled The only confirmed population of this species, a
abaxially and often with a narrow groove adjacent few hundred trees, is that at Mt. Panié. It is the most
to the keel; apex strongly incurved, acute. Stomata inaccessible of all species due to its location on the
few near the abaxial leaf base; adaxial stomata in summit of the highest mountain in New Caledonia,
parallel rows from base to apex, indistinct. Leaves to which a primitive, very steep footpath through
on adventitious branches acicular-linear, slightly dense rain and cloud forest gives the only access.
incurved, 7–18 × 1–1.7 mm, keeled abaxially above There is no prospect of mining due to the geology
base, pungent. Pollen cones terminal, initially erect of this mountain massif. It occurs within an estab-
but pendulous and curved at anthesis, 2–5 cm × 7–11 lished nature reserve, for which there are expansion
mm. Microsporophylls imbricate, spreading at 45° plans and, in future, to include it with the coastal
from a stout rachis, apically free at anthesis; stalk reefs in a UN World Heritage Site (Henry Blaffart,
thin and weak, linear, 2 mm long; lamina triangular, pers. comm., 2005).
2–3 × 1.5–2.5 mm, terminating in an acuminate apex. IUCN: VU (D2)
Pollen sacs not observed. Seed cones terminal on
short, stout foliage branches, erect, ovoid-oblong to
Uses
broadly ovoid, 7–9 × 5–6 cm. Bracts flabellate, 2–2.5
× 2–2.5 cm including rounded, thin membranous The wood is not exploited and no other uses have
wings, distally thickened and prominently keeled, been recorded for this species.
ending in a rostrate, curved, 15 × 2.5 mm tip. Seed
scales not wider than the seed; ligule fragile, 2 mm
long. Seeds obovoid-oblong, ca. 20 × 8 mm, smooth, Araucaria scopulorum de Laub., Trav. Lab. Forest.
brown when ripe. Toulouse T. 1 (8, 5): 1. 1969. Typ: New Caledonia:
Grande Terre, Province Nord, Cap Bocage,
H. S. MacKee 18760 (holotype P). Pl. 7, Fig. 53, 54
Distribution
New Caledonia: Grande Terre, Province Nord, sum- Araucaria bernieri J. T. Buchholz var. pumilio Silba,
mit of Mt. Panié. J. Int. Conifer Preserv. Soc. 7 (1): 21. 2000.
 5.5–9 × 4.5–7.5 cm. Bracts flabellate, 2–3(–3.4) × 1.8–
Etymology
2.5 cm including rounded, thin membranous wings,
The species epithet comes from the Latin scopulo- distally thickened and transversely keeled, ending in
rum = of cliffs or crags. a 4–6 mm long, upturned, acute tip. Seed scales not
or slightly wider than the seed; ligule fragile, 1.5–2
mm long. Seeds cuneate, 17–20(–24) × 6–9(–10)
Vernacular names
mm, smooth, light brown when ripe.
No common names have been recorded for this spe-
cies.
Taxonomic notes
212 Small trees in the north (i.e. on Mt. Poum and Dôme
Description
de Tiébaghi), alluded to by de Laubenfels (1972)
Monoecious trees to 20 m tall, to 0.5 m d.b.h.; trunk and previously regarded as A. bernieri, led Silba
straight. Bark to 2 cm thick, exfoliating in horizontal to describe A. bernieri var. pumilio based on Mac-
scales; inner bark reddish; outer bark grey with near- Kee 4832A. All trees from both localities that were
ly white patches. Resin exudate clear or yellow, dry- previously considered to be A. bernieri belong to
ing white or grey. Crown in mature trees irregular, A. ­scopulorum.
open, branched along greater part of trunk, domed
to flat-topped in older trees; branching according
Distribution
to Massart’s model. Primary first order branches of
mature trees in pseudo-whorls of 5–7, 1–2.5 m long, New Caledonia: Grande Terre, in two coastal areas,
spreading nearly horizontally to ascending, cadu- one along the central east coast and the other on the
cous. Adventitious foliage branches similar to pri- extreme NW coast (Mt. Poum, Dôme de Tiébaghi).
mary branches on mature trees but shorter. Foliage TDWG codes: 60 NWC
branchlets ascending or nearly erect near ends of
primary branches, forming two pectinate rows, 10–
Ecology
20 cm long and shortening towards the end, 4–9 mm
wide (including leaves), changing thickness along a Araucaria scopulorum occurs on rocky slopes and
single branchlet due to variable leaf sizes, flexible but ridges, often wind-swept, on serpentine or perido-
standing nearly erect. On mature trees leaves of two tite, in degraded forest and in ‘maquis minier’. Its
kinds: adult leaves on primary branches and semi-­ altitudinal range is 5–600 m a.s.l. and it is predomi-
juvenile leaves on adventitious branches. Adult leaves nantly coastal in its distribution. Especially typical
on ultimate branchlets ovate or obovate, 2.5–5.5(–6) of steep rocky slopes facing the sea in maquis or on
× 2.5–3.5 mm, keeled abaxially; apex incurved, sub­ nearly bare eroded rock and exclusively on ultra­
acute; leaves subtending pollen cones spreading, mafic substrates.
straight, narrowly oblong-lanceolate, narrower than
lower leaves on same branchlet and 2–3 times longer.
Conservation
Stomata mainly in 1–2(–3) rows nearest the keel on
each abaxial face, extending ± to apex, on the adaxial The distribution of this species nearly coincides with
face in ca. 20 rows from near base to apex. Semi- nickel mining areas and many populations are in or
juvenile leaves narrowly obovate to elliptic, 3.5–8 × adjacent to active nickel mines. When compared
1.2–2.3 mm, curved and with an abaxial keel. Pol- with the distribution map given by the Laubenfels
len cones terminal, solitary, initially straight but (1972) recent distribution data seem to register an
curved at anthesis, cylindrical, 2.5–5.5 cm × 10–12 increase, but this is an artefact due to more com-
mm. Micro­sporophylls spreading at 45° from a stout plete data, including collections made since 1972.
rachis; stalk thin and weak, 2–3 mm long; lamina In reality, numbers of trees and probably numbers
ovate-triangular, appearing rhombic when con- of (sub-)populations are declining. Fire and erosion
nate, 2.5 × 2 mm; apex acute. Pollen sacs 6–8, linear, have also been noted as threats. None of the known
straight, 3 mm long. Seed cones terminal on short, populations of this species are within a protected
stout foliage branches which widen below the cone, area at present.
usually solitary, erect, broadly ovoid to subglobose, IUCN: EN [B1ab(iii)+2ab(iii)]
 4

213

5
2

6
7

Pl ate 7. Araucaria scopulorum. 1. Habit of tree. 2. Foliage branch with pollen cones. 3. Section of
branchlet with leaves. 4. Pollen cone. 5. Microsporophyll with pollen sacs. 6. Seed cone. 7. Seed cone scales.
 at anthesis, 5–10 cm × 12–14 mm. Microsporophylls
Uses
imbricate, spreading at 45° from a stout rachis; stalk
The wood is not exploited and no other uses are re- thin and weak, 5 mm long; lamina more or less tri-
corded in New Caledonia. This species is in cultiva- angular, 3–4 × 2–2.5 mm; apex acute or acuminate.
tion in Tasmania. Pollen sacs ca. 10, linear, 2.5–3 mm long. Seed cones
terminal on short, stout foliage branches which
widen below the cone, solitary, erect, subglobose or
Araucaria subulata Vieill., Ann. Sci. Nat. Bot., broadly ovoid, 7.5–12 × 6.5–10 cm. Bracts flabellate,
sér. 4, 16: 55. 1862. Type: New Caledonia: Grande 2.5–3 × 2.2–2.7 cm including rounded, thin mem-
Terre, Province Sud, Canala, [“Pleine de Canala”], branous wings, distally thickened but thin-edged,
214 E. Vieillard 1278 (holotype P). ending in a caudate, upward curved, 8–10 mm long
tip. Seed scales not wider than the seed; ligule frag-
ile, 1–1.5 mm long. Seeds cuneate-oblong, 17–20 ×
Etymology
8–9 mm, striated longitudinally or smooth, light
The species epithet is from the Latin subulata = awl- brown when ripe.
shaped, in allusion to the leaf shape.
Distribution
Vernacular names
New Caledonia: Grande Terre, Province Sud.
No common names have been recorded for this spe- TDWG codes: 60 NWC
cies.
Ecology
Description
Araucaria subulata is emergent in dense humid sub-
Monoecious trees to 50 m tall, to 1.5 m d.b.h.; trunk montane to montane rainforest, at altitudes between
straight. Bark to 3 cm thick, exfoliating in horizontal 150 m and 1070 m a.s.l. It grows sometimes on cliffs;
flakes and strips, scaly; inner bark red-brown; outer apparently always on ultramafic soils or rubble de-
bark grey. Resin exudate white, very viscous. Crown rived from peridotite or serpentine. It occurs fre-
in mature trees columnar but open, branched along quently in ravines and steep valleys on the rainward
greater part of trunk, often widest in top; branching side (E and SE) of mountains, with accumulation of
according to Massart’s model. Primary first order litter and humus. This species is often co-dominant
branches of mature trees in pseudo-whorls of 5–7 with Agathis lanceolata and Montrouziera cauliflora
(–10) mostly restricted to the top of the tree, 0.5–1.5 (Guttiferae) and emerges from a 12–30 m tall canopy
m long, spreading or ascending, soon caducous. of numerous evergreen angiosperms, among which
Adventitious foliage branches numerous. Foliage sometimes Nothofagus sp. is abundant. Podocarpus
branchlets assurgent near ends of primary branches, sylvestris is another frequent coniferous tree in this
of more or less equal length (30–50 cm) in two pec- forest type. Tree ferns (Cyathea sp.) and palms are
tinate rows and shortening towards the end, 4.5–8 also common, the former especially in clearings.
mm wide (including leaves), remaining of ± equal
width from base to apex, flexible and whip-like. On
Conservation
mature trees leaves of two kinds: adult leaves on pri-
mary branches and semi-juvenile leaves on adventi- This species is relatively common in the south,
tious branches. Adult leaves on ultimate branchlets where it is associated with (remnants of) sub-mon-
subulate to broadly lanceolate, (3–)4–6 × 1–2.5(–2.7) tane rainforest. Outside protected areas, however,
mm, very glossy abaxially, strongly incurved, acute, this type of forest is often threatened by deforesta-
acuminate or obtuse. Stomata on the abaxial surface tion and fires from already deforested surrounding
near leaf base, on the adaxial surface in several rows areas. Mining is potentially of concern although at
for most of leaf length. Semi-juvenile leaves awl- present no mining operations are known to exist in
shaped or subulate, 4–8.5 × 0.5–1.2 mm; apex obtuse, any of its localities. Several populations are within
non-pungent. Pollen cones terminal, cylindrical, protected areas.
initially erect and straight, pendulous and curved IUCN: NT
 Uses
The wood is of high quality and has been locally ex-
ploited for use in carpentry.

215
 Athrotaxis D. Don, Ann. Nat. Hist. 1: 234. 1838. Arthrotaxis Endl., Gen. Pl. Suppl. 1:
1372. 1841, orth. var. Type: Athrotaxis selaginoides D. Don (Cupressaceae).

Greek: athroos = crowded; taxis = arrangement; re- Athrotaxis cupressoides D. Don, Ann. Nat. Hist. 1:
ferring to the leaves or seed cone scales. 235. 1838. Type: Australia: Tasmania, [“VDLand”
(Lindley’s handwriting)], R. C. Gunn [365] s.n.
(holotype CGE). Fig. 55
Description
Evergreen, monoecious trees to 30 m tall. Bark on
Etymology
216 young trees and branches smooth, flaking, on old
trees furrowed, exfoliating in thin flakes and long, The species epithet alludes to a resemblance with
shredding strips, reddish to light brown. Branches Cupressus.
forming a conical crown, sometimes layering to form
satellite trees. Leaves helically arranged in ranks of 5,
Vernacular names
decurrent, imbricate, small and appressed or longer
than 6 mm and spreading slightly or abruptly, rhom- Pencil Pine
bic-ovate to linear-lanceolate, keeled, lustrous green;
margins entire; apex obtuse or acute. Pollen cones
Description
at base surrounded by shortened leaves, solitary,
small; microsporophylls helically arranged, 10–15, Trees to 10–15 m, rarely 20 m tall; monopodial, erect,
broadly or narrowly triangular, bearing 2–4 oblong or very low branching to multi-stemmed; trunk up
pollen sacs; pollen subspherical without a papilla. to 0.5–1 m diam. Bark on old trees furrowed, ex-
Seed cones terminal on foliage branchlets, solitary, foliating in thin flakes and long, shredding strips,
globose or subglobose when opened, with to 30 light brown. Branches thick, ascending or spread-
helically arranged bract-scale complexes. Bract- ing, forming a conical crown, in old trees irregular
scale complexes clavate-peltate, thin or thick woody, and open, in multistemmed trees the lower branches
with a (prominently) exserted, incurved bract apex, form assurgent stems and separate crowns, layering
adaxially without visible seed marks. Ovules in a is common in peaty, saturated soil. Foliage branches
single row adaxial on the narrowed base of each numerous, mostly alternate, spreading, scaly, angu-
bract, anatropous. Seeds to 60 per cone, with two lar with keeled leaves, ultimate branchlets 3–4 mm
slightly unequal wings. Seedlings with 2 cotyledons. thick, mostly persistent. Leaves helically arranged in
ranks of 5, 3–6 × 2–3 mm (larger on leading, old-
3 species. er branchlets), fused at base, decurrent, imbricate,
appressed, oblong, the visible part rhombic-ovate,
keeled; margins thin hyaline, entire; apex obtuse,
Distribution
usually lustrous yellowish green to green; amphi­
Australia: Tasmania. stomatic, with widely spaced stomata (on the adax-
ial face in two faint bands). Pollen cones terminal
or subterminal, at base surrounded by shortened
Key to the species of Athrotaxis
leaves, 3–5 mm long, yellowish, turning brown.
1a. Leaves scale-like, 3–6 × 2–3 mm. Seed cone Micro­sporophylls helically arranged, 10–15, broadly
scales with thickened, centrally depressed apex triangular, obtuse, concave; upper margin denticu-
and small, protruding bract tip A. cupressoides late; short pedicellate; pollen sacs (2–)3–4, oblong.
1b. Leaves lanceolate to linear-lanceolate, (4–)7–18 Seed cones terminal on branchlets with normal
× 2–4 mm. Seed cone scales with thin, not de- scale leaves, solitary; mature cones globose or sub-
pressed apex and large, extended bract tip 2 globose, 10–16 × 10–14 mm; with 10–15 helically ar-
2a. Leaves 4–12 × 2–3 mm, mostly connate but with ranged bract-scale complexes which slightly part at
free apex A. ×laxifolia maturity. Bract-scale complex clavate-peltate, when
2b. Leaves 7–18 × 3–4 mm, the distal part spread- fully grown thick woody, with centrally depressed
ing A. selaginoides apex with a protruding, recurved, acute bract tip;
abaxial surface rough and wrinkled, reddish brown i­ mportance is in horticulture, where it is sometimes
to dark brown; adaxial surface striated, dark brown offered in the trade, but more often encountered in
without visible seed marks. Seeds 20–30 per cone, arboreta and botanic gardens.
obovate-oblong, 1.5–2 mm long, with 2 wings of
slightly unequal size and shape and ca. 1 mm wide
positioned more or less in one plane. Athrotaxis ×laxifolia Hook., Icon. Pl., n.s., 2: t. 573.
1843. Type: Australia: Tasmania, R. C. Gunn [369]
s.n. (holotype K).
Distribution
Australia: Tasmania, mainly on the Central Plateau,
Etymology
the Great Western Tier and westward mountains, 217
more scattered in the S of the island. The species epithet refers to the ‘lax’ or easily bend-
TDWG codes: 50 TAS able foliage.

Ecology Vernacular names

Commonly in the subalpine zone, where it of- Not recorded.
ten forms more or less dense stands, more scat-
tered in the transition zone to montane Eucalyptus
Description
woodland; at higher altitudes often on lake shores
or alongside streams and bordering ‘Buttongrass Trees to 10–15 m tall; monopodial, erect, in old trees
moorland’ or dwarf scrub. The altitudinal range is up to 0.5 m diam. Bark on old trees furrowed, exfo-
(700–)900–1300 m a.s.l. Associated with Sphagnum liating in thin flakes and long, shredding strips, light
cristatum, Gleichenia alpina, Leucopogon hookeri, brown. Branches thick, ascending or spreading,
Restio australis, Gymnoschoenus sphaerocephalus, forming a conical, dense crown, in old trees irregu-
Orites acicularis, Richea scoparia, R. pandanifolia, lar and open. Foliage branches numerous, mostly
Nothofagus gunnii, Astelia alpina, Diselma archeri, alternate, spreading, scaly, rough with free leaf api-
Pherosphaera hookeriana, Athrotaxis selaginoides, ces, ultimate branchlets slender, of unequal length,
and Eucalyptus coccifera; at lower altitudes with Eu- mostly persistent. Leaves helically arranged in ranks
calyptus spp., Nothofagus cunninghamii, and Phyllo- of 5, 4–12 × 2–3 mm (larger on leading, older branch-
cladus aspleniifolius. Soils are acidic, peaty or rocky lets), decurrent, imbricate, the distal part spreading
and usually wet. slightly, lanceolate, widest at the point of curvature,
keeled; margins entire; apex incurved, obtuse, usu-
ally lustrous light green; leaves amphi­stomatic, with
Conservation
few stomata proximally on the abaxial face and two
This species is extremely fire sensitive and much of bands of widely spaced stomata adaxially, some-
the total population was fire-killed mainly in wide- times epistomatic. Pollen cones terminal or sub-
spread fires on the Central Plateau in 1960/61. All terminal, at base surrounded by shortened leaves,
remaining areas of the species are now within IUCN 3–5 mm long, up to 4 mm wide, yellowish, turning
Category I-IV reserves. Regeneration of the species brown Microsporophylls helically arranged, ca. 15,
is limited by grazing, mainly by introduced sheep narrowly triangular, curved, concave; upper margin
and rabbits, of which the latter remain a problem. denticulate, short pedicellate; pollen sacs 2, oblong.
Dieback in which a species of the oomycete genus Seed cones terminal on branchlets with non-mod-
Phytophthora may be implicated occurs at some ified leaves, solitary; mature cones globose or sub-
high altitude sites. globose, 15–26 × 14–20 mm, with 14–18 helically
IUCN: VU (A 1a+c) arranged bract-scale complexes which part widely at
maturity. Bract-scale complex clavate-peltate, when
fully grown thin woody, with slightly depressed
Uses
apex and prominently extended, incurved, keeled
Usually a gnarled tree in its native habitat, this bract apex; abaxial surface rough and wrinkled, red-
species never was a valuable timber tree. Its main dish brown; adaxial surface striated, reddish brown
 ­ ithout visible seed marks. Seeds 30–40 per cone,
w Athrotaxis selaginoides D. Don, Ann. Nat. Hist. 1:
obovate-oblong, ca. 2 mm long, with 2 narrow wings 235. 1838. Type: Australia: Tasmania, [“VDLand”
of slightly unequal size and shape and 0.5–1 mm (Lindley’s handwriting)], R. C. Gunn [368] s.n.
wide positioned more or less in one plane. (holotype CGE). Fig. 56, 57

Taxonomic notes Etymology

In Tasmania, foresters and several botanists regard The species epithet alludes to similarity of the foli-
this species as a natural hybrid (F1) between the two age with the lycopod genus Selaginella.
other species. Flora of Australia 48 (1998) excluded
218 it with this argument. Farjon (2005a) discussed the
Vernacular names
evidence and observed that it was inconclusive. This
taxon may be of hybrid origin, as some other conifer King William Pine, King Billy Pine
species probably are, but it ‘breeds true’ and it is here
maintained as a nothospecies.
Description
Trees to 20–30 m tall; monopodial, erect, in old trees
Distribution
up to 1.5 m diam., with a straight, clear bole in dense
Australia: Tasmania, more scattered than the other forest but stunted in exposed sites. Bark on on old
two species, but sympatric with these. trees shallowly furrowed, exfoliating in thin flakes
TDWG codes: 50 TAS and long, shredding strips, reddish to light brown.
Branches thick, ascending or spreading, forming
a conical, dense crown, in old trees irregular and
Ecology
open. Foliage branches numerous, mostly alter-
Commonly growing with A. cupressoides and/or nate, spreading, rough with spreading, stiff leaves,
A. selaginoides as single or scattered individuals. ultimate branchlets thick, of unequal length, mostly
persistent. Leaves helically arranged in ranks of 5,
7–18 × 3–4 mm (larger on leading, older branch-
Conservation
lets), decurrent, the distal part spreading abruptly,
Because it occurs usually as solitary individuals, linear-lanceolate, widest at the point of curvature,
rarely in small groups, and generally with or near keeled; margins entire; apex incurved, acute, usu-
one or both of the other species A. cupressoides and ally lustrous light green; leaves amphistomatic, with
A. selaginoides, it is at risk of extinction mainly be- few stomata proximally on the abaxial face and two
cause of its low number of individuals overall. Near- broad and conspicuous bands of stomata separat-
ly all of these are now in reserves, where fire is the ed by a midrib but uniting distally on the adaxial
main threat. face, sometimes epistomatic. Pollen cones terminal
IUCN: VU D1 or subterminal, at base surrounded by shortened
leaves, 4–5 mm long, up to 5 mm wide, yellowish,
turning brown. Microsporophylls helically arranged,
Uses
ca. 15, narrowly triangular, curved, concave, short
This taxon has been taken into cultivation more of- pedicellate; upper margin denticulate; pollen sacs
ten than its apparent rarity in the wild suggests and 2, oblong. Seed cones terminal on branchlets with
can be seen in several arboreta, especially, but not non-modified leaves, solitary; mature cones glo-
exclusively, in the British Isles. bose or subglobose, 15–25(–30) × 15–20(–30) mm,
with 20–30 helically arranged bract-scale complexes scoparium, Eucalyptus spp., Diselma archeri, Phe-
which part widely at maturity. Bract-scale complex rosphaera hookeriana, and Phyllocladus aspleniifo-
clavate-spathulate, when fully grown thin woody, lius. Soils are more or less acidic, rocky (talus) or
with incurved apex and prominently extended, in- peaty and usually wet.
curved, keeled bract apex; abaxial surface rough and
wrinkled, dark brown; adaxial surface striated, dark
Conservation
brown without visible seed marks. Seeds 40–60 per
cone, obovate, 2–3 mm long, with 2 narrow wings of The major cause of past decline has been fire, with
slightly unequal size and shape and 0.5–1 mm wide about 1/3 of its habitat burnt in the 20th century. Like
positioned more or less in one plane. the other two species, A. selaginoides is sensitive to
fire. Another cause of decline has been logging. Al- 219
though 84% of forests are now in protected areas,
Distribution
fires still are a potential hazard. Tasmanian Govern-
Australia: Tasmania, widely distributed in the hig- ment policy precludes logging of this species in and
lands of W and SW parts of the island. outside these reserves.
TDWG codes: 50 TAS IUCN: VU (A 1a+c).

Ecology Uses
Commonly in the transition zone to (above) mon- King William pine provides some excellent wood for
tane Eucalyptus woodland; at higher altitudes less special uses like wood carving and turning. Howev-
frequent, there often on lake shores or alongside er, virtually all trees, including dead wood, are now
streams in sheltered places. The altitudinal range within protected areas and removal is prohibited. In
is (400–)730–1120 m a.s.l. Associated with Richea horticulture it is present only in arboreta and other
scoparia, R. pandanifolia, Nothofagus cunninghamii, botanical collections, especially in the British Isles,
N. gunnii, Atherosperma spp., Gahnia grandis, Telo- where several large trees can now be seen.
pea spp., Orites spp., Melaleuca spp., Leptospermum
 Austrocedrus Florin & Boutelje, Acta Horti Berg. 17 (2): 28. 1954. Type: Austrocedrus
chilensis (D. Don) Pic. Serm. & Bizzarri [Thuja chilensis D. Don] (Cupressaceae).

Latin: australis = southern; cedrus = cedar; therefore obtuse; lateral leaves decurrent, bilaterally flattened,
‘southern cedar’. 3–4.5 mm long on ultimate branchlets, incurved;
apex free or appressed, obtuse; margins entire; leaves
(nearly) hypostomatic; stomata in depressed green-
Description
ish white bands bordered by thick green margins
See the species description. on the abaxial side and in 2 small groups near base
220 on the adaxial side; transitional leaf form on leading
shoots (whip shoots) long decurrent (to 15 mm); lat-
Distribution
erals with a spreading, curved distal part terminat-
As for the species. ing in an acute apex. Pollen cones terminal, solitary,
5–7 × 2–3 mm; microsporophylls 12–20, decussately
arranged, imbricate, subpeltate with acuminate
Austrocedrus chilensis (D. Don) Pic. Serm. & Biz- apex; pollen spherical, in 3–4 abaxial pollen sacs
zarri, Webbia 32 (2): 482. 1978. Thuja chilensis near the lower margin of the microsporophyll. Seed
D. Don, in Lambert, Descr. Pinus, ed. 8°, 2: 128. cones terminal, often numerous on ultimate branch-
1832; Libocedrus chilensis (D. Don) Endl., Syn. Con- lets with non-modified leaves, maturing in one year
if.: 44. 1847. Type: Chile: [“Habitat in Andium Regni to ovoid-oblong, apically slightly flattened cones
Chilensis montibus”], J. A. Pavón y Jiménez [ex herb. 7–14 × 4–7 mm. Bract-scale complexes 4, decussate,
A. B. Lambert] s.n. (lectotype K). Fig. 58, 59 in pairs of very unequal size, the upper scales fertile
and twice as large as the lower, ovoid-oblong, with
a subapical curved umbo (bract tip), opening late,
Etymology
becoming woody, longitudinally striated or grooved
The species epithet refers to its country of origin. abaxially, smooth adaxially, brown. Columella coni-
cal to more or less clavate, 2–3 mm long. Seeds 2–4
per cone, angular-ovoid, ca. 4 × 2.5 mm, brown with
Vernacular names
light yellowish hilum and 2 wings, one rudimentary,
Chilean cedar, Cordilleran cypress; ciprès de la Cor- the other well developed, angular-oblong, 6–8 × 3–4
dillera, Ciprès (Spanish) mm.

Description Taxonomic notes

Shrubs or trees to 15–20(–25) m tall, evergreen, dio- First described as a species of Thuja, this taxon was
ecious, rarely monoecious; trunk multistemmed transferred to Libocedrus by Endlicher (1847), then
or monopodial to 1.5 m d.b.h. Bark smooth, soon to a monospecific genus Austrocedrus by Florin &
flaking, orange-brown turning grey with brown Boutelje (1954), who in creating the genus failed to
inner bark, fissued and scaly, exfoliating in long cite the correct basionym. This error was corrected
strips. Branches numerous, spreading, assurgent in by Pichi Sermolli & Bizzarri, who thereby validated
higher part of crown, persistent, forming a conical the currently used combination as cited above.
or pyramidal or more rounded crown in trees. Foli-
age branches dense, spreading in horizontal planes
Distribution
or assurgent, nearly erect in young trees, ultimate
branchlets plagiotropic, opposite or nearly so, short, Central and S Chile: Biobio, La Araucania, Los
directed forward, covered by leaves, persistent, turn- Lagos, Maule, O’Higgins, Santiago; S Argentina:
ing orange- to red-brown, soon grey. Leaves scale- Chubut, Neuquén, Rio Negro.
like, decussate, in opposite ranks, dimorphic; facial TDWG codes: 85 AGS-CB AGS-NE AGS-RN CLC-BI
leaves very small, ca. 1 mm, partly hidden by larger CLC-LA CLC-MA CLC-OH CLC-SA CLS-LL
laterals, incurved-appressed, keeled, acuminate or
 Ecology Conservation
This species has a natural distribution accross near- This species, though still wide spread, is under vari-
ly 12 degrees of latitude in the Andes, from 32° to ous pressures that make it vulnerable to extinction.
42–44° S, thus ranging from a Mediterranean type Felling, expansion of human habitation and agricul-
climate to the cold montane Patagonian steppe cli- ture, transformation of natural forest to plantation
mate. It generally shifts from N to S from the west- forest with exotic trees (Eucalyptus, Pinus), and fires
ern slopes to the eastern slopes of the Andes, leaving have all played a role in the decline of its area of occu-
more mesic conditions to other tree species. It of- pation. A dieback problem has been reported in the
ten occurs in pure stands or mixed with Nothofagus Argentinian populations (Havrylenko et al., 1989).
dombeyi after quickly covering former disturbance Seed viability is often low (down to less than 5%) due 221
areas (fire, earthquakes, volcanic deposits) but also to insect predation (M. Gardner, pers. comm., 1996).
grows associated with Fitzroya ­cupressoides in the Only a relatively small proportion of its populations
coastal ranges and with Nothofagus spp. in the An- is within protected areas in both countries (ca. 15%
des. Austrocedrus chilensis gradually disappears in in Argentina). Grazing of livestock and feral deer is
the wetter regions to the south where Notho­fagus a problem both outside and inside these reserves.
will eventually dominate after disturbances. Its IUCN: VU [A2 c; B 2 a, b (iii)].
tolerance to shade is relatively low, but apparently
somewhat higher than of its common associate
Uses
Nothofagus dombeyi, so that some individuals can
persist in the understorey. It is longer-lived than its Timber of Chilean cypress is used for general car-
competitor, though not reaching more than 1000 pentry and furniture. Its fragrant and durable qual-
years, usually around 500 years, so it can fill gaps left ity makes it especially suitable for panelling and for
by fallen Notho­fagus dombeyi. Its edaphic require- outdoor furniture. It is a useful species for horticul-
ments are lower than for most tree species in the ture, as in cultivation it usually grows a fastigiate,
region, so it can occupy sandstone, volcanic, or gra- columnar habit (an as yet unexplained phenom-
nitic outcrops where other species will not establish, enon seen also in other members of Cupressaceae,
thereby extending its range into the wetter latitudes e.g. Calocedrus decurrens). Despite this it is still un-
of the Southern Andes. Where it borders the Patago- common and largely confined to arboreta and oth-
nian steppe it is the only tree species, but does not er botanical collections. A few cultivars have been
attain large size there. mentioned in the 19th century but may no longer be
available.
 Austrotaxus R. H. Compton, J. Linn. Soc., Bot. 45: 427. 1922. Type: Austrotaxus
spicata R. H. Compton (Taxaceae).

Latin: australis = southern; taxus = yew; therefore dark green above, light green below; margins
‘southern yew’. slightly revolute. Midrib forming a narrow groove
on the adaxial (upper) face from base to apex, on
the abaxial side raised, 0.8–1.2 mm wide. Stomata
Description
in two broad bands of numerous wavy, intermittent
See the species description. lines on tha abaxial side only, bordered by narrow
222 green margins and the midrib. Pollen cones axil-
lary, solitary but aggregated near base of new foli-
Distribution
age shoots, on a scaly short peduncle or dwarf shoot,
As for the species. elongating to 12–15 mm, catkin-like, creamy white,
microsporophylls12–18, remote, spirally arranged,
reniform, concave, on the adaxial side bearing two
Austrotaxus spicata R. H. Compton, J. Linn. Soc., or three rows of globose pollen sacs, up to 8 sacs in
Bot. 45: 427. 1922. Type: New Caledonia: Grande the outer row and 2–3 larger ones in the inner row.
Terre, Province Nord, Mt. Ignambi, R. H. Compton Seed-bearing structures axillary, solitary near base
1155 (holotype BM). Fig. 60 of new foliage shoots, on a very short, scaly pedun-
cle or dwarf shoot, the upper scales enlarged, im-
bricate, triangular, enclosing a single apical ovule.
Etymology
Aril almost entirely enclosing the seed, ovoid and
The species epithet (Latin spicatus = spike) refers to slightly flattened when mature, 12–15 mm long, 7–9
the pollen cones. mm wide, smooth, glaucous green ripening to or-
ange, only leaving the seed apex free. Seed proper
ovoid-oblong, yellowish brown with a darker apex
Vernacular names
and minute micropylar tip.
New Caledonia yew.
Distribution
Description
New Caledonia: Grande Terre.
Trees to 25 m tall; trunk erect and straight to 60 cm TDWG codes: 60 NWC
d.b.h. in forest. Bark fibrous, exfoliating in short
strips or on the lower trunk becoming tesselated,
Ecology
dark reddish brown weathering grey. Branches nu-
merous, forming a dense crown. Foliage branchlets Austrotaxus spicata is locally common in the mon-
subopposite or more or less verticillate, spreading at tane tropical rainforests of the northern two thirds
45–70° from the main branch, retaining leaves only of the main island, Grande Terre. It has been found
towards ends, terete, with obtriangular to oval leaf at altitudes between 500 m and 1350 m a.s.l. and ap-
scars. Terminal buds globose, ca. 3 mm wide and pears on serpentine (ultramafic soils) as well as on
2–3 mm long, with triangular, apiculate scales free acidic mica schist (Mt. Canala, Mt. Panié) or gneiss
at apex. Leaves alternate (helical), on juvenile plants (Mt. Ignambi). It is associated with other conifers
longer than on mature trees, up to 17 cm long; on and angiosperms commonly found in the south of
mature plants (4–)6–9(–12) cm long, spreading for- the island, so its (seeming?) absence there is some-
ward at 30–60° to shoot axis, crowded towards ends what mysterious. Its superficial resemblance to
of branchlets except on vigorous leading shoots, Podo­carpus spp., with which it may occur together,
narrowly lanceolate, 3.5–7(–9) mm wide near the makes it difficult to detect unless the tree is fertile,
middle, straight or slightly falcate, gradually wid- but other things being equal this applies to its entire
ening from a 5–8 mm long petiolate base, gradu- range.
ally tapering to an acute apex; coriaceous, ­lustrous
 montane serpentine deposits (garnierite) within its
Conservation
altitudinal range.
This interesting species, an endemic to Grande IUCN: NT
Terre, appears to be relatively common and is not
specifically exploited for its timber. It occurs often
Uses
in inaccessible places and at least in several forest
reserves, e.g. Table Unio, Mt. Panié, and Roches No uses have been recorded of this species and it is
de Ouaième (proposed). The quest for anti-cancer not known to be in cultivation except as a few young
drugs found in the alkaloid compounds of the fam- plants in perhaps three botanic gardens (BGCI da-
ily Taxaceae, which led to the exploitation of several tabase, accessed May 2008). Its wood properties
species, does not seem to have affected this isolated presumably resemble those of other members of 223
species seriously. On the other hand, deforestation Taxaceae, which are not commercial timber trees
probably has had an impact on it, particularly in but can be put to specialized uses like furniture,
nickel mining concessions in the north of the island, wood turning and arts and crafts. This interesting
and it is not safe from further developments in this species should be grown more in botanic gardens
increasingly spreading activity, particularly aimed at and other tree collections where climatic conditions
are suitable.
 Callitris Vent., Decas Gen. Nov.: 10. 1808. Frenela Mirb., Mém. Mus. Hist. Nat.
13: 30. 1825. [Frenela triquetra Spach] Type: Callitris rhomboidea R. Br. ex Rich. &
A. Rich. (Cupressaceae).

Octoclinis F. Muell., Trans. & Proc. Philos. Inst. Vic-

Distribution
toria 2: 21. 1858. Type: Octoclinis macleayana F. Muell.
[Callitris macleayana (F. Muell.) F. Muell.]. Nothocal- Australia including Tasmania; New Caledonia.
litris A. V. Bobrov & Melikyan, Komarovia 4: 84.
2006. Type: Nothocallitris sulcata (Parl.) A. V. Bo-
Synopsis
224 brov & Melikyan [Callitris sulcata (Parl.) Schltr.].
In the Monograph of Cupressaceae and Sciadopitys
Greek: calli = beautiful. (Farjon, 2005a) two sections were accepted: section
Callitris with 14 species and section Octoclinis with a
single species, Callitris macleayana. A phylogenetic
Description
analysis based on multiple-gene DNA sequence data
Shrubs or trees, evergreen, monoecious. Resin cavi- and comprehensive taxon sampling is still wanting.
ties in leaves. Bark on trunks of large trees thick, Despite this, the separation of the morphologically
deeply fissured, hard (soft and stringy in one spe- and ecologically distinctive species C. macleayana at
cies). Branches spreading or ascending, forming a section rank seems warranted and is here accepted.
bushy, conical, pyramidal or broadly domed crown.
Fastigiate forms common in several species in na- Callitris sect. Callitris
ture. Ultimate branchlets covered with leaves, lateral Species: Callitris baileyi C. T. White, C. canes-
branchlets shed continuously. Leaves scale-like, in cens (Parl.) S. T. Blake, C. columellaris F. Muell.,
alternate whorls of 3 (in one species also with ac- C. drummondii ((Parl.) F. Muell., C. endli-
icular, spreading leaves in whorls of 4), decurrent, cheri (Parl.) F. M. Bailey, C. monticola J. Gar-
appressed, linear, abaxially convex or keeled, con- den, C. muelleri (Parl.) Benth. & Hook. f. ex
nate but parting with the thickening of branches, F. ­Muell., C. neocaledonica Dummer, C. oblon-
persisting several years, extremely variable in size ga Rich. & A. Rich., C. preissii Miq., C. rhom-
dependent on growth of branches but on ultimate boidea R. Br. ex Rich. & A. Rich., C. roei
branchlets mostly less than 5 mm long; margins usu- (Endl.) F. Muell., C. sulcata (Parl.) ­Schltr.,
ally minutely denticulate, apical part slightly differ- C. verrucosa (A. Cunn. ex Endl.) F. ­Muell.
entiated (e.g. thicker) from main length of the leaf.
Pollen cones solitary or with 2–3 together at the tips Callitris sect. Octoclinis (F. Muell.) Benth. &
of branchlets, small, short cylindrical; microsporo- Hook. f.
phylls 8–20 in whorls of 3 or decussate, with 2–6 Species: Callitris macleayana (F. Muell.)
abaxial pollen sacs. Seed cones terminal on short F. ­Muell.
leafy shoots, solitary or more often clustered and
serotinous, (sub)globose when closed, early decidu-
Key to the sections of the genus Callitris
ous or more often persistent on branches and stems.
Bract-scale complexes in two alternate whorls of 3 1a. Seed cones with two whorls of ternate scales,
(in one species also in two whorls of 4), with the up- only on branches with adult scale leaves. Bark
per whorl usually the largest, valvate, thick, smooth, smooth or hard and scaly Section Callitris
rugose or with verrucae on the abaxial surface and 1b. Seed cones with two whorls of ternate as well as
with light seed marks on the adaxial surface; bract quadrate scales, the latter on branches with ju-
tips subapical, small. Columella distinct, variably venile needle leaves, both occurring in mature
shaped, often trimerous, the lobes alternating with trees. Bark soft and stringy Section Octoclinis,
the upper whorl of scales. Seeds moderately numer- a single species Callitris macleayana
ous, with 2–3 wings. Seedlings with 2 cotyledons.

15 species.
 C. muelleri
Key to the species of section Callitris
11a. Seed cones mostly longer than wide, with mark-
1a. Seed cones at the terminal end of thin foliage edly different lengths of scales in each whorl
branchlets, solitary or a few together, caducous C. oblonga
2 11b. Seed cones angular-globose; scales more equal
1b. Seed cones on short, thickened foliage branch- in length 12
lets, usually clustered, persistent 6 12a. Seed cones whitish grey to metallic grey, weath-
2a. Dorsal surface of adult leaf rounded, not ering to dull grey. Restricted to SW Western
keeled 3 Australia C. roei
2b. Dorsal surface of adult leaf distinctly (obtusely 12b. Seed cones darker coloured. Eastern Australian
or acutely) keeled 4 species 13 225
3a. Shrub or small tree. Seed cone scales narrowly 13a. Dwarf tree 1–3(–4) m tall; bark remaining
oblong and nearly equal in size, ca. 3 mm wide. smooth; leaves on ultimate branchlets 2–3 mm
Endemic in New Caledonia C. sulcata long. Restricted to a small area in NE New
3b. Tree. Seed cone scales unequal in size, wider South Wales and SE Queensland C. monticola
than 4 mm. Widespread in mainland Australia 13b. Shrub or tree to 10–15 m; bark soon scaly, fis-
C. columellaris sured; leaves on ultimate branchlets 3–6 mm
4a. Shrub or small tree. Juvenile and transitional long. Widespread from Queensland to Tasma-
leaves persisting on young trees. Endemic to nia C. rhomboidea
New Caledonia C. neocaledonica
4b. Trees. Juvenile and transitional leaves restricted
to seedlings. Native in mainland Australia 5 Callitris baileyi C. T. White, Proc. Linn. Soc.
5a. Bark of lower trunk thick, deeply fissured; New South Wales 48: 449. 1923. Type: Australia:
leaves obtusely keeled C. endlicheri Queensland, Blackbutt Range, Benarkin,
5b. Bark on lower trunk thin, shallowly fissured; W. D. Francis [BRI 192911] s.n. (lectotype BRI).
leaves acutely keeled C. baileyi
6a. Dorsal surface of adult leaf rounded, not
Etymology
keeled 7
6b. Dorsal surface of adult leaf distinctly (obtusely This species was named after Frederick M. Bailey
or acutely) keeled 9 (1827–1915) who collected the first specimens.
7a. Seed cones verrucose when mature, more or
less covered with small or large verrucae 8
Vernacular names
7b. Seed cones smooth or rugose when mature,
verrucae absent C. canescens Bailey’s Cypress-pine
8a. Verrucae variable in density, often quite large in
proportion to the cone, sometimes nearly ab-
Description
sent C. preissii
8b. Verrucae dense, equally distributed and never Trees to 15–18 m tall, often less than 10 m; mono­
nearly absent, small in proportion to the cone podial, usually branching low and often fastigiate, up
C. verrucosa to 50 cm d.b.h. Bark on trunk thin, shallowly fur-
9a. Mature seed cones globose, smooth, with very rowed or fissured, hard, slowly exfoliating in small
small subapical bract tip not deforming the flakes and strips, greyish, inner layers light brown.
cone scale 10 Branches ascending at ca. 45° or steeper, slender,
9b. Mature seed cones angular-globose or longer forming a conical or pyramidal, often symmetri-
than wide, with large subapical bract tip de- cal crown. Foliage branchlets sparse, ascending or
forming the cone scale 11 erect, twisted and rigid, triangular in cross section,
10a. Seed cones (10–)12–15 mm wide, with tapering 0.8–1.2 mm diam., covered with closely appressed
base from thickened shoot terminus; leaves ob- leaves. Leaves in alternate whorls of 3, decurrent,
tusely keeled C. drummondii closely appressed, apices appressed or free on some
10b. Seed cones 15–25 mm wide, with abrupt base leading shoots (whip shoots), connate but parting
from shoot terminus; leaves acutely keeled with thickening of branches, persisting several years,
 l­inear, 2–5(–7) mm long on ultimate branchlets, of fires. Several populations are now within pro-
0.7–1.2 mm wide, abaxially acutely keeled; margins tected areas (National Parks), but decline is likely to
denticulate near obtuse or acute apex; epistomatic, continue outside these localities.
stomata in two narrow, marginal bands; abaxial IUCN: VU (A1c)
surface smooth, green. Pollen cones solitary or 2–3
together and terminal on ultimate branchlets, ob-
Uses
long, 2–3 × 1–1.2 mm, yellowish green turning light
brown; microsporophylls 8–14, in alternate whorls of The wood of this species was traditionally used for
3 or decussate near apex, peltate, with hyaline, erose- fence posts. It is extremely rare in cultivation, lim-
denticulate margins, bearing 2–3 abaxial pollen sacs ited to a few botanic gardens.
226 near the lower margin. Seed cones terminal on short,
thin, leafy shoots, solitary, maturing in 1–1.5 years,
caducous, subglobose to broadly ovoid when closed, Callitris canescens (Parl.) S. T. Blake, Proc. Roy.
10–13 × 9–12 mm, smooth but becoming rugose Soc. Queensland 70: 39. 1959. Frenela canescens
when open, brown or purplish brown weathering Parl. in Candolle, Prodr. 16 (2): 448. 1868. Type:
grey. Bract-scale complexes in 2 alternate whorls of Australia: Western Australia, [“Int. S. W. Australia”],
3, deeply incised towards cone base but not opening J. S. Roe s.n. (lectotype G?, n.v., isolectotype K). Fig.
very wide; upper ones largest, ca. 10 × 5 mm, tapering 61
to an obtuse apex; smaller ones narrowly triangular,
acute; bracts largely included, showing a prominent
Etymology
apex below the apex of each scale; adaxial surface
reddish brown or brown with light seed marks. The species epithet means ‘greyish’ or ‘turning grey’
Columella robust in comparison to small cone (as and may refer to the seed cones.
high as shortest scales in open cones), 3–4 × 2–2.5
mm, more or less triangular in cross-­section, usually
Vernacular names
slightly narrowed at base, obtuse. Seeds 2–4(–5) on
each scale, ovoid or triangular, flattened, up to 6–7 Morrison’s Cypress-pine
mm including wings (seed body 2–3 mm), brown
with large, whitish concave hilum, wings 2, lateral,
Description
very unequal in size and shape, up to 4 mm long.
Shrubs or small trees to 5–6 m; trunk short, usually
branching low or monopodial. Bark on stems wrin-
Distribution
kled, finally scaly near base, exfoliating in flakes,
Australia: SE Queensland (Blackbutt Range, Bunya grey. Branches numerous, ascending or nearly erect
Mts., Coast Range, Leichhardt Highway, West More- (fastigiate), contorted, higher order branches slen-
ton), NE New South Wales (Clarence River, Killar- der, terete, forming a conical, pyramidal or bushy
ney, Tabulam). dense crown. Foliage branchlets numerous, erect,
TDWG codes: 50 NSW-NS QLD-QU ultimately very slender, terete, 0.6–0.7 mm diam.,
covered with closely appressed leaves, persistent.
Leaves in alternate whorls of 3, decurrent, closely
Ecology
appressed; apices barely free, connate but parting
In Eucalyptus forest or woodland, with Eucalyptus with thickening of branches; persisting 3–4 years,
crebra or other species, often in small groups scat- linear, 1.5–3 mm long on ultimate branchlets, up to
tered through forest; in hills and low mountains on 1 mm wide, abaxially convex; margins denticulate
basalt and volcanic ash or in sandy soil with boulders. near apex; epistomatic, stomata near the margins of
The climate is mild, with most rainfall in the summer. leaves only; abaxial surface granulate with globular
epidermal cells. Pollen cones terminal on ultimate
branchlets, numerous, ovoid or subglobose, 1.5–2
Conservation
× 1 mm, yellowish green turning brown; micro-
Decline has mainly come from changes in habitat sporophylls 9–12, in alternate whorls of 3, peltate,
due to pressures from grazing and increased hazard with erose margins, bearing 3(–4) abaxial pollen
sacs near the lower margin. Seed cones terminal on tle interest in the native flora and prefer to import
short, thickened, leafy shoots, solitary or more com- shrubs and trees cultivated in Britain. In Mediter-
monly aggregated, persistent, depressed globose ranean countries (and perhaps with a warming cli-
when closed, (10–)12–14(–16) × (11–)13–15(–19) mm, mate soon elsewhere in Europe?) this species should
smooth to coarsely rugose when dry, light brown be cultivated more often.
turning (metallic) grey. Bract-scale complexes in 2
alternate whorls of 3, the upper ones largest, 10–14
× 6–8 mm, tapering to an obtuse apex; the smaller Callitris columellaris F. Muell., Fragm. 5: 198.
ones narrower 8–10 × 5–7 mm, acute; bracts almost 1866. Type: Australia: New South Wales,
entirely included, showing a minute prickle below Richmond River, [“Richmond River Entrance”],
the apex of each scale; adaxial surfaces dark grey J. A. Henderson [ex herb. F. von Mueller] MEL 227
with whitish seed marks towards base. Columella 1596576 (lectotype MEL). Fig. 62
simple, 2–3 mm long, obtuse or acute-triangular.
Seeds 3–6 on each scale, of variable shapes and sizes Frenela robusta A. Cunn. ex Endl. var. microcarpa
up to 7 mm including wings (seed body 1–3 mm), Benth., Fl. Austral. 6: 237. 1873; Callitris columellaris
blackish brown with light hilum, wings 2–3 sur- F. Muell. var. microcarpa (Benth.) Govaerts, World
rounding the seed as a 2–3 mm wide margin, curved Checklist Seed Plants 3 (1): 12. 1999.
to accomodate neighbouring seeds when growing. Callitris intratropica R. T. Baker & H. G. Smith,
Res. Pines Austral.: 172. 1910; Callitris columellaris
F. ­Muell. var. intratropica (R. T. Baker & H. G. Smith)
Distribution
Silba, Phytologia Mem. 7: 16. 1984.
SW and SE Western Australia, South Australia (Eyre Callitris columellaris F. Muell. var. campestris Silba,
Peninsula, Murray River, Nullarbor Plain, Yorke Phytologia Mem. 7: 16. 1984.
Pen­insula). Callitris glaucophylla J. Thompson & L. A. S. John-
TDWG codes: 50 SOA WAU-WA son, Telopea 2 (6): 731. 1986.

Ecology Etymology
In dwarf scrubland (‘kwongan’) with Myrtaceae The species epithet refers to a ‘columella’, i.e. the cen-
(Melaleuca), Proteaceae (e.g. Banksia, Dryandra, tral, columnar structure in the seed cone.
Grevillea, Hakea), or low Eucalyptus woodland
(‘mallee’) with other Myrtaceae (e.g. Melaleuca),
Vernacular names
Proteaceae (e.g. Banksia, Grevillea, Hakea), Aca-
cia spp., and Allocasuarina spp.; often in strips of White Cypress-pine, White Pine, Murray River
natural vegetation between roads and ploughed Pine, Murray Pine, Western Sand Cypress, Western
fields or in disturbed vegetation; on plains or low Cypress, Cypress Pine, Northern Cypress-pine
ridges (‘breakaways’) in leached white, yellow or
red (gravelly) sand or loam, or on laterite (hardpan)
Description
over granite, sandstone or limestone. The altitudi-
nal range is from 250 m to 380 m a.s.l. The climate Trees to 20 m tall; trunk monopodial, usually
is characterized by warm, dry summers and winter branching low or occasionally forked, up to 50 cm
rainfall. d.b.h. Bark soon scaly, on trunk deeply fissured,
hard, exfoliating in small flakes and strips, grey-
brown. Branches spreading or near the top ascend-
Conservation
ing, long, forming a conical, pyramidal or broadly
IUCN: LC domed, more or less open crown. Foliage branch-
lets numerous, spreading or ascending, ultimately
very slender, terete, 0.7–1.0 mm diam., covered
Uses
with closely appressed leaves, persistent. Leaves in
This species is not in cultivation outside botanical alternate whorls of 3, decurrent, closely appressed,
collections. Australian gardeners generally show lit- apices appressed or free on some leading shoots
 (whip shoots), connate but parting with thicken- TDWG codes: 50 NSW-NS NTA QLD-QU SOA VIC
ing of branches, persisting several years, linear, 2–6 WAU-WA
mm long on ultimate branchlets, 0.4–0.8 mm wide,
abaxially convex; margins denticulate near slightly
Ecology
broadened acute-acuminate apex; epistomatic, sto-
mata in two marginal lines, abaxial stomata near In a wide range of semi-arid to more mesic habitats,
base of leaves only or absent; abaxial surface weakly from banks or dunes near the sea to rocky canyon
verrucose, green or glaucous, variable with many bottoms and sandstone plateaux in the interior. In
intermediate shades of colour. Pollen cones numer- open Eucalyptus forest or low woodland (‘mallee’)
ous, solitary or paired and terminal on ultimate with Eucalyptus spp., Melaleuca spp., Allocasua-
228 branchlets, ovoid-oblong, 3–4 × 1.5 mm, ­yellowish rina spp., Acacia spp., Banksia spp., Grevillea spp.,
green turning orange-brown; microsporophylls Triodia spp., and Callitris endlicheri; in disturbed
10–18, in alternate whorls of 3, peltate, with erose (grazed) open woodland; in the dry interior in Aca-
margins, bearing 3(–4) abaxial pollen sacs near the cia low woodland or scrub (‘mulga’) with A. aneura
lower margin. Seed cones terminal on short, thin, and other Acacia spp., Eremophila spp., in hum-
leafy shoots, solitary but often numerous on higher mock grassland/ scrub with ‘spinifex’ (Triodia sp.);
order branches, maturing in 1–1.5 years, caducous, and in rocky (sandstone) canyons and on ridges
globose when closed, (7–) 10–18 × (10–)12–20 mm, with Eucalyptus spp., Acacia spp., and Macrozamia
smooth or rugose to slightly pustulate, glaucous macdonnellii. On (coastal) sand dunes near sea level,
brown or purplish brown weathering grey. Bract- on plains, hills, tablelands, ridges and low moun-
scale complexes in 2 alternate whorls of 3, deeply tains to 1300 m a.s.l. in white, yellow, brown or red
incised towards cone base when opened; upper ones sand or loam or on gravelly laterite (hardpan) over
largest, 8–12 × 6–8 mm, tapering to an obtuse apex; sandstone, ironstone, conglomerate, gypsum, shale,
smaller ones narrower 6–8 × 4–6 mm, acute; bracts basalt or granite. The climate varies from mesic and
almost entirely included, showing a minute prickle mild on the coast of New South Wales to arid in the
below the apex of each scale; adaxial surfaces red- interior of Australia; for most of its range this spe-
dish brown or dark brown with light seed marks to- cies occurs within the zone with summer rainfall.
wards base. Columella variably shaped, up to 6 mm
long, obtuse or acute-triangular. Seeds 4–10 on each
Conservation
scale (the higer number on larger scales), triangular,
flattened, up to 8 mm including wings (seed body IUCN: LC
up to 4–5 × 2–3 mm), dark brown with whitish con-
cave hilum; wings 2 on opposite sides, 4–5 mm wide,
Uses
more or less equal in shape and size.
Timber of this species was (is?) used for panelling in
cabinet making, and also for fence posts when these
Taxonomic notes
were still predominantly of wood. As with many
Treatment of Callitris glaucophylla and C. intra­ species in Cupressaceae, the wood is resistant to
tropica as taxonomic synonyms under a (slightly) rot and, in this case, also to termite attack. Baker &
broader circumscription of C. columellaris has been Smith (1910) reported a decline in its availability in
explained and justified in detail in the Monograph many regions without efforts to restock it. Its exploi-
of Cupressaceae and Sciadopitys (Farjon, 2005a). tation is now much less intensive and regeneration
is taking place in many localities. Its horticultural
use is very limited. In Australia, there seems to be
Distribution
little interest in this and other native conifers, but
Australia: In all states and territories except Tas- its variability in foliage colour from dark green to
mania, but absent from low lying deserts and moist extremely glaucous should afford scope for horticul-
coastal regions. tural selection and planting in regions with a Medi-
terranean or even desert climate.
Callitris drummondii (Parl.) F. Muell., Syst. Census incised towards cone base and hence not opening
Austral. Pl. 1: 109. 1882. Frenela drummondii Parl. widely; upper ones largest, 10–12 × 8–10 mm, taper-
in Seemann, J. Bot. 1: 35. 1863. Type: Australia: ing to an angular-obtuse apex; smaller ones nar-
Western Australia, Southern Coastal Region, [“In rower 8–10 × 6–8 mm, acute, all very thick; bract tip
Nova Hollandia austro-occidentali ad Cynorem minute, only visible in smooth, closed cones below
flumen legit Cl. J Drummond”], J. Drummond [ex the apex of each scale; adaxial surfaces dark brown
herb. F. von Mueller] 433 (holotype FI). with light seed marks at margins near base. Colu­
mella very thick, 3–5 mm long and wide at base,
(tri-)angular or terete, obtuse. Seeds usually only
Etymology
2 on each scale, angular, flattened, up to 7 mm in-
This species was named after J. Drummond, who cluding wings (seed body 2–3 × 2 mm), brown with 229
collected the type specimens. whitish concave hilum; wings 2 on opposite sides,
1–2 and 2–4 mm wide, unequal in shape and size.
Vernacular names
Distribution
Drummond’s Cypress-pine
SW Western Australia, in a strip along or near the
coast of the Southern Ocean from Albany to Cape
Description
Arid.
Shrubs or small trees to 5–6(–10) m; multistemmed TDWG codes: 50 WAU-WA
or monopodial, usually branching low, up to 20 cm
d.b.h. Bark smooth on young trees and branches,
Ecology
soon scaly, on the lower stem(s) fissured, hard, grey.
Branches spreading to ascending, forming a conical In scrubland (‘Melaleuca-thicket’) or low open Eu-
or broad, rounded crown. Foliage branchlets nu- calyptus woodland (‘mallee’) with Eucalyptus spp.,
merous, ascending, ultimately relatively coarse, ar- Melaleuca spp., and Acacia spp.; also along margins
ticulate and triangular in cross section, 1.5–2.2 mm of (dry) salt lakes and flats, in dry stream beds and
diam., covered with closely appressed leaves, per- in disturbed vegetation on roadsides; on soils vary-
sistent. Leaves in alternate whorls of 3, decurrent, ing from white to dark brown sand, loam, silt or clay
closely appressed, connate but parting with thicken- over sandstone, limestone or gravelly laterite (hard-
ing of branches, persisting several years, linear, 2–6 pan). Its altitude ranges from 10 m to 250 m a.s.l. The
mm long on ultimate branchlets, 1–1.8 mm wide, climate is characterized by warm, dry summers and
abaxially keeled; margins denticulate near acute or winter rainfall.
obtuse apex; epistomatic, stomata in two marginal
lines; abaxial surface weakly verrucose or smooth,
Conservation
yellowish green or green. Pollen cones solitary or
in pairs and terminal on ultimate branchlets, ovoid- Outside reserves this species occupies mainly strips
oblong, 3–6 × 1.5–2 mm, yellowish green turning of (semi)natural vegetation left alongside roads, a
light brown; microsporophylls 12–20, proximal ones situation which has made the species more vulner-
in alternate whorls of 3, distal ones decussate, sub- able to fires in these areas as these now occur more
peltate, with hyaline-denticulate margins; apex acu- frequently. Overgrazing is also a threat in some
minate; bearing 3 abaxial pollen sacs near the lower ­areas. On the other hand, several good size popula-
margin. Seed cones terminal on short, 5–6 mm thick, tions are within reserves where these factors can be
leafy shoots, solitary but often grouped along lim- managed to benefit the natural vegetation.
ited lengths of stems, maturing in 1–1.5 years, per- IUCN: VU (A1c)
sistent, (depressed-)globose when closed, tapering
at base to thickened terminus of shoot, (10–)12–15
Uses
× (10–)12–17 mm, smooth or rugose when dry with
opened scales, purplish grey to metallic grey. Bract- This species is not in cultivation outside botani-
scale complexes in 2 alternate whorls of 3, not deeply cal collections. In Mediterranean countries (and
 ­ erhaps with a warming climate soon elsewhere in
p ­ argin. Seed cones terminal on short, thin, leafy
m
Europe?) this species should be cultivated more of- shoots, solitary but often grouped together in loose
ten since it grows into an attractive shrub. clusters, maturing in 1–1.5 years, caducous, ovoid-
globose when closed, 13–20 × 10–16 mm, smooth or
finely rugose when open, brown or purplish brown
Callitris endlicheri (Parl.) F. M. Bailey, Syn. weathering grey. Bract-scale complexes in 2 alter-
Queens­land Fl.: 497. 1883. Frenela endlicheri nate whorls of 3, deeply incised towards cone base
Parl., in Candolle, Prodr. 16 (2): 449. 1868. Type: when opened; the upper ones largest, 10–16 × 8–12
Australia: Victoria, Murray Valley District, Warby mm, tapering to an obtuse apex; the smaller ones
Ranges, [“Futters Range”], F. von Mueller MEL narrower 8–10 × 5–7 mm, acute; bracts almost en-
230 503743 (lectotype MEL). tirely included, showing a minute prickle below the
apex of each scale; adaxial surface reddish brown or
dark purplish brown with light seed marks. Colu-
Etymology
mella trilobed to tripartite, 2–8 mm wide, obtuse.
This species was named after Stephan F. L. Endlicher Seeds 5–9 on each scale, ovoid or triangular, flat-
(1804–1849) an Austrian botanist who published an tened, up to 8 mm including wings (seed body ca. 3
early manual of conifers. × 2 mm), dark brown with large, yellowish concave
hilum; wings 2–3 on sides, 2–3 mm wide, more or
less equal in shape and size.
Vernacular names
Black Cypress-pine, Black Pine, Red Cypress-pine,
Distribution
Red Cypress, Red Pine, Mountain Pine, Scrub Pine,
Murray Pine Australia: New South Wales, Queensland, E Victo-
ria.
TDWG codes: 50 NSW-NS QLD-QU VIC
Description
Trees to 15–20 m tall; monopodial, usually branch-
Ecology
ing low and occasionally fastigiate, up to 50 cm
d.b.h. Bark soon scaly, on lower trunk very thick, In open Eucalyptus woodland or forest, in second-
deeply fissured, hard, dark grey-brown, inner layers ary scrub or woodland, or in grassland, often in
red-brown. Branches ascending usually at ca. 45°, disturbed or altered vegetation with invasives like
short or long in fastigiate crowns, slender, forming a Cytisus scoparia and Pinus radiata. Associated with
conical or pyramidal, more or less open crown. Fo- Eucalyptus spp., other Myrtaceae (e.g. Callistemon,
liage branchlets numerous, spreading or ascending, Kunzea, Leptospermum), Proteaceae (e.g. Banksia,
ultimately very slender, triangular in cross section, Grevillea), Acacia spp., Allocasuarina sp., Bursaria
0.6–1 mm diam., covered with closely appressed lasiophylla, Daviesia sp., Dodonaea viscosa, and Cal-
leaves, persistent. Leaves in alternate whorls of 3, litris rhomboidea. On usually very shallow (gravelly)
decurrent, closely appressed, apices appressed or sand or loam, or in deep rock crevices, or amongst
free on some leading shoots (whip shoots), connate boulders over sandstone, conglomerate, slate,
but parting with thickening of branches, persist- quartzite or granite, often on steep slopes; also in
ing several years, linear, 2–6 mm long on ultimate disturbed soil on roadsides. The altitude varies be-
branchlets, 0.3–0.7 mm wide, abaxially keeled; mar- tween 350 m and 1100 m a.s.l. The climate is mesic
gins denticulate near slightly broadened acute-acu- to summer dry; frost and snow occur occasionally at
minate apex; epistomatic, stomata in two narrow, higher altitudes.
marginal bands; abaxial surface weakly verrucose,
(dark) green. Pollen cones numerous, solitary or
Conservation
paired and terminal on ultimate branchlets, ovoid-
globose to oblong, 1.5–2.5 × 1–2 mm, yellowish green Cutting for timber and fence posts has had an impact
turning orange-brown; microsporophylls 8–14, in in the past, but these pressures are now substantial-
alternate whorls of 3, peltate, with denticulate mar- ly alleviated by an economic shift to other sources
gins, bearing 2–4 abaxial pollen sacs near the lower for these materials. The species is not ­substantially
a­ ffected by conversion of its habitat to agriculture covered with decurrent leaves, persistent. Leaves of
or plantation forestry due to its occurrence on poor two types on mature trees, scale leaves in alternate
sites. whorls of 3, acicular leaves mostly in whorls of 4,
IUCN: LC all decurrent. Scale leaves predominantly in upper
parts of crown, closely appressed, apices appressed
or free on some leading shoots (whip shoots),
Uses
connate but parting with thickening of branches,
Timber of this species was much used for fence posts persisting several years, linear, 1.5–3 mm long on
in the past, the worked wood also for construction, ultimate branchlets, 0.6–0.8 mm wide, abaxially
and when finely figured it was and still is valued for strongly keeled; margins erose-hyaline near acute
indoor finishing of rooms (panelling, wain­scoting), apex. Acicular (juvenile) leaves predominantly in 231
furniture and some decorative applications. The lower parts of crown, free part spreading, 8–15 mm
bark was used for tanning and some resin was long, abaxially keeled; margins entire; apex pun-
tapped from the tree trunks. It is not considered at- gent. Leaves epistomatic, stomata in two marginal
tractive as an amenity tree in Australia, but some bands; abaxial surface smooth, green. Pollen cones
planting of stony wasteland with this undemanding solitary and terminal on ultimate branchlets with
tree has been successfully undertaken. It is present scale leaves, oblong, 4–6 × 2–2.5 mm (elongating
in a few botanic gardens in countries with a mild, to ca. 8 mm), yellowish green turning light brown;
summer-dry climate. microsporophylls 15–20, in alternate whorls of 3,
peltate, with erose margins and acute apex, bear-
ing 3–4 abaxial pollen sacs near the lower margin.
Callitris macleayana (F. Muell.) F. Muell., Rep. Seed cones terminal on short, thick, recurved, leafy
Burdekin Exped.: 17. 1860. Octoclinis macleayana and often glaucous shoots, mostly solitary, maturing
F. Muell., Trans. & Proc. Philos. Inst. Victoria 2: in 1 year, persistent, ovoid-conical or conical when
22. 1858. Type: Australia: New South Wales, Port closed, 15–35 × 18–35 mm, smooth becoming rugose,
Macquarie, Tacking Point, W. S. Macleay s.n. maturing dark brown. Bract-scale complexes in 2
(holotype K). Fig. 63, 64 alternate whorls of 3 on branches with scale leaves
and of 4 on branches with acicular leaves, deeply in-
cised towards cone base but not opening wide, very
Etymology
thick near base, nearly equal in size and shape, con-
This species was named after William John Macleay cave abaxially, tapering to an acute, slightly recurved
(1820–1891) parlementarian of New South Wales, apex; adaxial surfaces angular, reddish brown with
Australia. light seed marks at the base. Columella short, tri-
lobed or 4-lobed, often reduced to low ridges. Seeds
not exceeding 10–12 per cone, ovoid, flattened, up to
Vernacular names
8 mm long, dark brown with whitish concave hilum;
Stringybark Pine, Port Maquarie Pine, Brush Cy- only one wing develops to a length of 8–10 mm, the
press-pine other rudimentary.

Description Taxonomic notes

Trees to 30(–39) m tall; monopodial, usually self- This species is unique in the genus in having poly-
pruning forming a clear bole to 10 m or more, up morphism in mature trees in phyllotaxis and leaf
to 1 m d.b.h. Bark soon scaly, on trunk deeply fis- shape. Small scale leaves occur in alternating whorls
sured, soft, stringy and fibrous, sometimes scaly, of three and are appressed, as in other species.
exfoliating in long strips, reddish brown or light ­Acicular leaves are arranged in alternating whorls
brown. Branches spreading wide or ascending, long of four and have spreading free parts. Because the
and thick, forming a pyramidal or broadly domed, number and arrangement of cone scales strictly fol-
more or less open crown. Foliage branchlets in tufts, low shoot phyllotaxis and both types of shoot are
spreading, ultimately slender, triangular in cross- fertile, both 6- and 8-scale cones occur, often on the
section if with scale leaves and ca. 1 mm diam., same tree. Four-whorled phyllotaxis is restricted to
 the ­seedlings in other species of Callitris and does
Uses
not reappear in the crown of mature trees. The only
real distinction then is the persistence of juvenile The timber of this species is free of knots in large
traits in the foliage of this species; the difference boles and has been used for shingles and weather-
in the number of cone scales follows from it and is boards, and also for cabinet making and joinery. As
not an independent character. The different bark in many species of Cupressaceae it is decay resistant
structure, reminiscent of that found on Sequoia and and, especially relevant in Queensland, termite re-
­Sequoiadendron, may be a more distinct feature. As sistant as well, so it has been used for fence posts.
an adaptive trait it is likely to be a parallelism, but The use of metal rods for agricultural fencing has
a more detailed anatomical investigation or phylo- largely replaced this application. Stringybark Pine
232 genetic analysis would be necessary to confirm or is not known in horticulture outside a few botanic
refute this hypothesis. gardens.

Distribution
Callitris monticola J. Garden, Contr. New South
Australia: On the eastern slopes of the Great Divid- Wales Natl. Herb. 2 (5): 385. 1957. Type: Australia:
ing Range and along the coast in New South Wales Queensland, [“Wallangarra, Queensland”],
N of Newcastle to extreme SE Queensland. W. A. W. de Beuzeville NSW 22815 (holotype NSW).
TDWG codes: 50 NSW-NS QLD-QU Pl. 8

Ecology Etymology
In subtropical rainforest remnants with Ficus The Latin species epithet monticola means ‘living in
obliqua, Drypetes australasia, Archontophoenix mountains’.
cunning­hamiana, and Podocarpus elatus; also in wet
sclerophyll forest bordering rainforest with Eucalyp-
Vernacular names
tus microcorys, E. pilularis, Ceratopetalum apetalum,
Allocasuarina torulosa, Acacia spp., Tristania spp., Steelhead
and Trochocarpa laurina. In hollows, gullies, shel-
tered near-coastal flats, or rain-facing (E) slopes,
Description
on red-brown clay (dolorite) or loam. The altidudi-
nal range is from near sea level to around 1000 m Dwarf trees 1–3(–4) m, sometimes a shrub, mono­
a.s.l. The climate is subtropical to warm temperate ecious; trunk monopodial or multistemmed,
and mesic, with a rainfall peak during the summer branching low, up to 10 cm d.b.h. Bark smooth to
(range 1000–2000 mm/a); frost does not occur in slightly scaly, thin, slowly exfoliating in small flakes,
the northern localities and is rare and light in the grey. Branches spreading or ascending at ca. 45°,
N. S. W. upland localities. long and pliant, forming an open, irregular or py-
ramidal crown. Foliage branchlets sparse, spreading
irregularly or more or less erect, arranged in more
Conservation
or less planated sprays towards ends of branches;
Large trees in more accessible terrain have in the ultimate branchlets very slender, triangular in cross
past been logged more extensively than they have section, 0.6–0.8 mm diam., covered with closely ap-
regrown to date, with the result that such trees have pressed leaves, persistent. Leaves in alternate whorls
become very rare. However, the species has regener- of 3, decurrent, closely appressed, apices appressed
ated well in most of its natural localities. Forest fires or some free especially on some leading shoots
are a potential threat that requires adequate man- (whip shoots), connate but parting with thicken-
agement with the conservation of this species as an ing of branches, persisting several years, linear, 2–3
objective. Its thick bark may make large trees with mm long on ultimate branchlets, 0.5–0.6 mm wide,
branch-free boles resistant to light fires that only abaxially prominently keeled; margins denticulate
burn through the undergrowth. near slightly broadened acute apex; epistomatic,
IUCN: NT
 233

1
4

5
6
7

Pl ate 8 . Callitris monticola. 1. Habit of trees. 2. Foliage branch. 3. Branchlet with juvenile and adult
leaves. 4. Pollen cone. 5. Shoot with pollen cones and seed cones. 6. Cluster of seed cones. 7. Seeds.
 s­ tomata mainly in two narrow marginal bands;
Conservation
abaxial surface densely verrucose, yellowish green to
glaucous. Pollen cones solitary and terminal on ulti- This species has a restricted distribution with very
mate branchlets, ovoid-globose, 2.5 × 2 mm, yellow- localized populations in a vegetation type that is un-
ish green turning orange-brown; microsporo­phylls der increased threat of fires. On one locality (sum-
9–12, in alternate whorls of 3, peltate, apiculate, with mit area of Bald Rock) it could not be found again
erose-denticulate margins, bearing 2–3 abaxial pol- despite a thorough search in 1997 by me and earlier
len sacs near the lower margin. Seed cones termi- searches by J. Williams from Armidale; only the
nal on short, thick, leafy shoots, mostly aggregated common C. rhomboidea remains there.
in dense, compact clusters on stems and branches, IUCN: VU (A1c)
234 maturing in 1–1.5 years, persistent, subglobose when
closed, 12–20 × 12–18 mm, verrucose or nearly
Uses
smooth, slightly rugose when open, dull brown,
weathering dull grey. Bract-scale complexes in 2 al- No uses are known of this species.
ternate whorls of 3, not deeply incised towards base
and opening only slightly; upper ones largest, 10–16
× 6–10 mm, with parallel or slightly widening sides Callitris muelleri (Parl.) Benth. & Hook. f. ex
and obtuse apex; smaller ones wider at base, triangu- F. Muell., Syst. Census Austral. Pl. 1: 109. 1882.
lar, with acute apex; bract tip forming a small, acute Frenela muelleri Parl., in Candolle, Prodr. 16 (2):
umbo below the apex of each scale; abaxial surface 450. 1868. Type: Australia: New South Wales, Syd­
convex, brown turning grey, with conspicuous white ney, Port Jackson, [“South Head”], A. Thozet [ex
(dotted) margins; adaxial surface dark brown with herb. F. von Mueller] MEL 503745 (lectotype MEL).
inconspicuous seed marks. Columella large, most- Fig. 65, 66
ly trilobed, up to 8 mm wide, obtuse. Seeds 3–5 on
­basal part of each scale, more or less triangular, flat-
Etymology
tened, up to 8 mm including wings (seed body 4–5
× 2–3 mm), dark brown, with concave hilum; wings This species was named after F. J. H. von Mueller
2 on each side, up to 4 mm wide, unequal or nearly (1825–1896), the famous German botanist working
equal in shape and size. on the Australian flora.

Distribution Vernacular names

Australia: SE Queensland, NE New South Wales, Illawarra Cypress-pine, Illawarra Pine
from ca. 28°S to ca. 30°S straddling the Great Divid-
ing Range.
Description
TDWG codes: 50 NSW-NS QLD-QU
Dwarfed trees or shrubs to 3–4.5(–6) m; mono­
podial or multistemmed, branching low and often
Ecology
fastigiate, up to 15 cm d.b.h. Bark smooth, not fis-
This species occurs in low Eucalyptus woodland or sured, thin, slowly exfoliating in small flakes and
forest and is virtually restricted to rock outcrops strips, grey. Branches ascending at ca. 45° or erect,
and cliff edges thinly covered with scrub. Associated short or long in fastigiate crowns, slender, forming
with Eucalyptus spp., Allocasuarina rigida, Xanthor- a conical to pyramidal, in fastigiate forms columnar
rhoea sp., Banksia spp., Leptospermum spp., Mirbelia crown. Foliage branchlets numerous, in dense tufts
confertifolia, and Restio stenocoleus; in very shallow towards end of main branches, ultimately slender,
soil amongst boulders or in crevices in granitic rock triangular in cross section, 0.7–1 mm diam., covered
(Adamellite), trachyte, or sandstone. The altitudinal with closely appressed leaves, persistent. Leaves in
range is from 560 m to 1360 m a.s.l. The climate is alternate whorls of 3, decurrent, closely appressed,
mild and mesic, with a peak of rainfall during the apices appressed or free on some leading shoots
summer. (whip shoots), connate but parting with thickening
of branches, persisting several years, linear, 2–8 mm m a.s.l. The climate is temperate and mesic and frost
long on ultimate branchlets, 0.5–0.7 mm wide, abax- can occur in winter at the higher altitudes.
ially keeled; margins denticulate near slightly broad-
ened obtuse or sometimes acute apex; epistomatic,
Conservation
stomata in two narrow, marginal bands; abaxial sur-
face minutely verrucose, green. Pollen cones nu- Some of the earlier populations reported from the
merous, solitary or with 2–3 together, terminal on Sydney (Port Jackson) area have undoubtedly been
ultimate branchlets, ovoid-globose to oblong, 2–3 × reduced in area of occupancy due to urbanisation.
1–1.5 mm, yellowish green turning orange-brown; Fires, if too frequent, will threaten regeneration
microsporophylls 9–12, in alternate whorls of 3, pel- success. Away from the urbanised areas the spe-
tate, with denticulate margins, bearing 2–3 abaxial cies, though not widespread, seems to hold its own 235
pollen sacs near the lower margin. Seed cones ter- in sites that support no commercial land use of any
minal on short, thick, leafy shoots, solitary but often kind.
grouped together in small clusters on branches and IUCN: LC
stems, maturing in 1–1.5 years, persistent, subglobose
or globose when closed, 15–25 × 15–25 mm, smooth
Uses
and lustrous when closed, coarsely rugose when
open, greenish or glaucous when growing, maturing There are no economic uses reported of this species.
to brown or purplish brown weathering grey. Bract-
scale complexes in 2 alternate whorls of 3, very thick,
not deeply incised towards cone base when opened; Callitris neocaledonica Dummer, J. Bot. 52: 239.
upper ones much larger than the lower, to 20 × 13 1914. Nothocallitris neocaledonica (Dummer)
mm, usually widest above the middle and tapering A. V. Bobrov & Melikyan, Komarovia 4: 86. 2006.
to an obtuse apex; smaller ones narrower and taper- Type: New Caledonia: Grande Terre, Province
ing to an acute apex; a minute prickle (bract tip) Sud, Ngoye River, [“auf den Bergen am Ngoye”],
below the apex of each scale; adaxial surface red- R. Schlechter 15179 (holotype K).
brown or dark brown with numerous whitish seed
marks. Columella triangular, sometimes flattened,
Etymology
ca. 3 mm long and wide, acute, dark brown to black.
Seeds 5–10 on each scale, ovoid or triangular, flat- The species epithet refers to its country of origin.
tened, up to 5 × 3 mm, dark brown with lighter col-
oured concave hilum; wings 2, 2–5 mm wide, more
Vernacular names
or less equal in shape and size.
Nié
Distribution
Description
Australia: E-central New South Wales, from areas
near the coast to the table lands of the Blue Mts. E of Shrubs or small trees 3–7(–10) m tall; trunk mono­
Sydney. podial or very short, branching low, up to 50 cm
TDWG codes: 50 NSW-NS diam. at base. Bark soon scaly, on lower trunk be-
coming shallowly fissured, rough, exfoliating in long
strips, brown weathering grey. Branches spread-
Ecology
ing and assurgent, thick and long, often contorted,
In scrub vegetation associated with Myrtaceae (e.g. forming a sympodial, rounded and open crown.
Calytrix, Eucalyptus apiculata, E. stricta, E. sieberi, Foliage branchlets in dense tufts at ends of main
Leptospermum), Proteaceae (e.g. Banksia, Grevillea), branches, assurgent or erect, ultimate branchlets
Allocasuarina distyla, Lomandra glauca, and Caustis short, rigid, thick, apprearing conspicuously articu-
pentandra. On ‘skeletal’ sandy or gravelly soil and late with scale leaves, triangular in cross section, 1.5–
among boulders or in rock crevices of sandstone on 2 mm diam., covered with closely appressed leaves,
cliffs or talus slopes, sometimes congregating on dirt green, persistent. Leaves of two types on plants up
road verges. The altitude ranges from 500 m to 1050 to 1 m tall. Mature (scale) leaves (and transitional
 leaves) in alternate whorls of 3, decurrent, closely reserves, but fire is a hazard and adequate manage-
appressed, apices appressed or some free especially ment of this increasing problem is urgently needed.
on some leading shoots (whip shoots), connate but IUCN: NT
parting with thickening of branches, linear, 2.5–5
mm long on ultimate branchlets, ca. 1 mm wide,
Uses
acutely keeled abaxially; margins erose-denticulate
near obtuse or acute apex; epistomatic, stomata in There are no commercial uses recorded of this spe-
two narrow marginal bands; abaxial surface smooth cies.
on keel, minutely verrucose on sides, green. Pollen
cones solitary and terminal on ultimate branchlets,
236 ovoid or subglobose, 2–3 × 2–2.5 mm, yellowish Callitris oblonga Rich. & A. Rich., in A. Richard
turning brown; microsporophylls 9–12, in alternate (ed.) Comm. Bot. Conif. Cycad.: 49. 1826. Type:
whorls of 3, peltate, with erose-denticulate margins, Australia: Tasmania, [“Nouvelle Hollande”],
obtuse-acuminate, bearing 3–4 abaxial pollen sacs J. L. C. Dumont d’Urville s.n. (holotype P).
near the lower margin. Seed cones few, terminal on
short, thin, scale-leaved shoots, solitary, maturing in Callitris oblonga Rich. & A. Rich. subsp. corangensis
1 year, caducous (or persistent a short time after seed K. D. Hill, Fl. Australia 48: 716. 1998.
dispersal), ovoid when closed, 8–10 × 6–7 mm, with Callitris oblonga Rich. & A. Rich. subsp. parva
convex scales when open, smooth, greenish matur- K. D. Hill, Fl. Australia 48: 717. 1998.
ing to brown. Bract-scale complexes in two alternate
whorls of 3, not opening very wide, nearly equal in
Etymology
size and shape, 7–8 × 3 mm, narrowly oblong, slight-
ly tapering, rectangular in cross-section, with most- The species epithet refers to the oblong (longer than
ly included bracts forming a prominent and acute wide) seed cones.
umbo below the apex of each scale; adaxial surface
smooth, red-brown with inconspicuous seed marks.
Vernacular names
Columella very long, 5–6 × 1.5–2 mm, angular, with
triangular apex. Seeds 1(–2) at base of each upper Tasmanian Cypress-pine, Dwarf Cypress-pine, Na-
scale, ovoid or triangular, flattened, 5–7 × 2–2.5 mm, tive Cypress, River Pine, Pigmy Cypress-pine
light brown, with small hilum; wings 2 on each side,
up to 7 × 1.5 mm, unequal in shape and size.
Description
Shrubs or small trees to 7–8 m tall, often less than
Distribution
4 m; monopodial or multistemed from a short basal
New Caledonia, Grande Terre, Province Sud. trunk, usually branching low and often fastigiate, up
TDWG codes: 60 NWC to 20 cm d.b.h. Bark soon scaly, on the lower trunk
of largest trees fissured, hard, slowly exfoliating in
small flakes and strips, grey-brown. Branches as-
Ecology
cending or erect, long in fastigiate crowns, slen-
In scrub vegetation (‘maquis minier’) or low sclero- der, forming a conical or pyramidal, open or dense
phyll forest on slopes or along creeks, in shallow soil crown, or a more or less rounded shrub in less fa-
over ultramafic (serpentine) rocks. The altitudinal vorable sites. Foliage branchlets ascending to erect,
range is between 560 m and 1500 m a.s.l. The climate ultimate branchlets triangular in cross section, 1–1.2
is low montane tropical with high rainfall through- mm diam., covered with closely appressed leaves,
out the year. persistent. Leaves in alternate whorls of 3, decurrent,
closely appressed, apices appressed or free on some
leading shoots (whip shoots), connate but part-
Conservation
ing with thickening of branches, persisting several
This species has a limited distribution in localized years, linear, 3–12 mm long on ultimate branchlets,
populations in the southern massifs. It is protected 0.7–1 mm wide, abaxially keeled; margins denticu-
in the Montagnes des Sources and Mt. Humboldt late near slightly broadened acute-acuminate apex;
epistomatic, stomata in two narrow, marginal bands; Acacia spp., Leptospermum spp., Allocasuarina spp.,
abaxial surface weakly verrucose, green or glaucous Bursaria spinosa, Callistemon paludosis, Jacksonia
green. Pollen cones subterminal in clusters of (2–)3– scoparia, Lomandra longifolia, Epacris microphylla,
5(–6) on ultimate branchlets, ovoid-globose, 1–1.5 × 1 Hakea spp., Lomatia myrtifolia, Pteridium sp., and
mm, yellowish green turning orange-brown; micro­ grasses (Poaceae). The altitudinal range is from 10 m
sporophylls 8–12, in alternate whorls of 3, peltate, to 1300 m a.s.l., with the Tasmanian populations
rounded, with entire margins, bearing 2–3 abaxial mostly below 100 m a.s.l. Soils are (gravelly) sand or
pollen sacs near the lower margin. Seed cones termi- loam, often rocky with granitic boulders or bedrock,
nal on very short, thick, leafy shoots, rarely solitary acidic, and sometimes silt or clay, subject to regu-
and often grouped together in compact clusters on lar flooding. In a few cases in Tasmania, C. oblonga
branches and stems, maturing in 1–1.5 years, persist- appears to occur away from streams on floodplains, 237
ent, ovoid-oblong or ovoid-utriculate when closed, invariably on disturbed sites grazed by cattle, and in
(12–)15–20(–25) × (10–)12–16(–20) mm, smooth, ru- New South Wales on moderate slopes with seepage
gose when dry, blackish brown or black, weathering water. The climate is temperate and mesic, without
grey. Bract-scale complexes in 2 alternate whorls of prolonged periods of drought, but with higher rain-
3, deeply incised towards cone base when opened, fall levels in the uplands of New South Wales than in
thick, parting only slightly; upper ones largest, up NE Tasmania.
to 20 × 10 mm, tapering to a (recurved) obtuse
apex; smaller ones 1/2 to 3/5 the size of the larger
Conservation
ones, acute or obtuse; adaxial surfaces dark purplish
brown with light seed marks. Columella small, tri- Briggs & Leigh (1988) list Callitris oblonga (N. S. W.)
angular to trilobed, 2–3 mm long, obtuse. Seeds 5–15 and “Callitris sp. 1” [= C. oblonga in Tasmania] as
on each scale, ovoid or triangular, flattened, up to Vulnerable (3VCa in CSIRO conservation coding).
10 mm including wings (seed body 3–4 × 2–3 mm), More recent research (Nadolny & Benson, 1993) has
dark brown to black with large, yellowish concave confirmed a wider occurrence in the New England
hilum; wings 2, lateral, 3–4 mm wide, more or less Range of N. S. W. than was previously known. Some
equal in shape and size. of the populations are in conservation areas, but
most are outside these on rangeland or bushland.
Threats are conversion to agriculture by clearance of
Taxonomic notes
natural vegetation (in particular Corang River and
For a discussion of the two subspecies described in Tasmania), grazing of livestock, associated invasive
Flora of Australia 48 (1998), here treated as taxo- species (especially Ulex europaeus) and fires if too
nomic synonyms of Callitris oblonga, the reader frequent (with intervals of 3 years or less). Some
is referred to the Monograph of Cupressaceae and populations are (now) very small and isolated. In
­Sciadopitys (Farjon, 2005a). Tasmania a recovery programme is under way using
several approaches.
IUCN: VU (A1c)
Distribution
Australia: New South Wales, in two disjunct areas:
Uses
in the New England Tablelands and adjacent moun-
tains, and along the Corang River, a tributary of the There is a limited ornamental use of this species in
Shoalhaven River; and in NE Tasmania along sev- Tasmania. In cultivation, as with many species in
eral rivers. Cupressaceae, Tasmanian Cypress-pine grows con-
TDWG codes: 50 NSW-NS TAS sistently in a fastigiate habit. In the wild, this habit
is commonly found in Tasmania but, as far as I have
seen, absent in New South Wales. It is possible that
Ecology
there is a genetic basis for this, but environmental
Mostly riparian, in Eucalyptus woods or shrubland; factors also seem to play a role, although it remains
in Tasmania often in disturbed vegetation with in- unclear what these are specifically. From what I have
vasives like Ulex europaeus, Rubus discolor, and seen in this species in both areas, it could be that
Crataegus sp. Associated taxa are Eucalyptus spp., more adverse conditions for fast growth prevent
 fastigiate branching and promote spreading branch- or acute apex; epistomatic, a few abaxial stomata
ing instead. The fastigiate form should be cultivated near base of leaves only or absent, adaxial stomata
more widely, as it forms a well shaped erect shrub or in two marginal bands and a few below apex; abax-
small tree. ial surface verrucose, yellowish green or some-
times glaucous green. Pollen cones solitary or 2–3
together, terminal on ultimate branchlets, ovoid-
Callitris preissii Miq., in Lehmann, Pl. Preiss. oblong, 3–5 × 1.5–2.5 mm, yellowish green turning
1: 643. 1845. Type: Australia: Western Australia, orange-brown; microsporophylls 15–20, in alternate
Rottnest Island, C. A. Gardner 175 (epitype whorls of 3 or decussate, peltate, ovate-orbicular,
PERTH), L. Preiss 1310 (lectotype L). Fig. 67 with entire or erose margins, bearing 2–4 abaxial
238 pollen sacs near the lower margin. Seed cones ter-
Callitris tuberculata R. Br. ex R. T. Baker & minal on short, thick, leafy shoots, solitary or more
H. G. Smith, Res. Pines Austral.: 99. 1910. often clustered on branches and stems, maturing in
Callitris preissii Miq. subsp. murrayensis J. Garden, 1–1.5 years, persistent, (depressed-)globose to ovoid-
Contr. New South Wales Natl. Herb. 2 (5): 373. 1957; globose when closed, (15–)20–30(–35) × (18–)22–35
Callitris preissii Miq. var. murrayensis (J. Garden) mm, nearly smooth to coarsely rugose and vari-
Silba, Phytologia Mem. 7: 17. 1984; Callitris gracilis ably covered with large verrucae, brown or purplish
R. T. Baker subsp. murrayensis (J. Garden) K. D. Hill, brown to blackish, weathering grey. Bract-scale
Fl. Australia 48: 716. 1998. complexes in 2 alternate whorls of 3, not deeply in-
cised towards cone base when opened, very thick;
the upper ones largest, 12–30 × 8–13 mm, tapering to
Etymology
an obtuse apex; the smaller ones narrower, acute or
This species was named after Ludovicus (Ludwig) obtuse; adaxial surfaces grey-brown or brown with
Preiss, who collected this species on Rottnest Island light seed marks. Columella short, thick, up to 5 mm
in 1839. long and as wide at base, obtuse or acute-triangu-
lar. Seeds (7–)9–12(–16) on each scale, triangular
to ovoid, flattened, up to 10 mm including wings
Vernacular names
(seed body up to 3–5 × 2–3 mm), dark brown or red-
Rottnest Island Pine, Mallee Pine, Murray Pine, brown with whitish concave hilum; wings 2(–3) on
Mountain Pine, Mountain Cypress-pine, Lachlan opposite sides, 2–4 mm wide, more or less equal or
Pine, Light Pine, White Pine, Common Cypress- unequal in shape and size.
pine
Taxonomic notes
Description
Flora of Australia 48 (1998) took a narrow view of
Shrubs or trees to 25 m tall; monopodial or mul- C. preissii and included only the populations on
tistemmed, usually branching low and frequently Rottnest Island and “around Perth” in it. The Flo-
forked, up to 1 m d.b.h. Bark soon scaly, on large ra accepted C. tuberculata as a widespread species
trunks thick, deeply fissured, stringy, exfoliating in occurring along the coast and in the interior of
large strips, light brown; outer bark grey. Branches Western Australia and C. gracilis, including subsp.
spreading or ascending, long, forming a pyramidal murrayensis, occurring in South Australia, Victoria
or broadly domed, more or less open crown, or a and New South Wales. Investigation of populations
pyramidal or rounded shrub. Foliage branchlets on sand dunes at Bremer Bay, Western Australia
spreading or ascending, ultimately very slender, (which according to Flora of Australia taxonomy
terete, 0.7–1.0 mm diam., covered with closely ap- belong to C. tuberculata), and populations in similar
pressed leaves, persistent. Leaves in alternate whorls habitat in two of the ‘classical’ localities of C. preis-
of 3, decurrent, closely appressed (except on some sii (Garden Island and Woodman Point) by me in
whip shoots), connate but parting with thickening 1997 revealed that they are the same. Multistemmed
of branches, persisting several years, linear, 1.5–5 shrubs are the dominant growth form at Woodman
mm long on ultimate branchlets, 0.5–1 mm wide, Point and common at Bremer Bay and on Garden
abaxially convex; margins denticulate near obtuse Island. The wartiness of cones as well as their size are
quite variable in all these populations. The plants in be extensive in the interior parts of the range) and
the interior are more distinct, especially those that winter rainfall.
grow on lateritic soil types, being (smaller) shrubs,
never a tree, and having smaller cones. In Victoria,
Conservation
I observed a similar situation: on sand dunes in
the Murray River Basin and in the ‘Little’ and ‘Big’ IUCN: LC
Deserts I found mostly trees, but in the hardpan
areas between the dunes I found shrubs. Cones are
Uses
less densely warty here than in Western Australia,
but considering its variation in that state, I think this The larger trees with monopodial growth that oc-
is insufficient evidence to separate C. gracilis from cur locally have been used for timber in local con- 239
C. preissii and a broad concept of the species is here struction, but its economic value is now limited and
retained. Garden (1957) further confounded the tax- no commercial harvest occurs. It is only known in
onomy of this species by postulating two naturally horticulture from arboreta and botanic gardens in
occurring hybrids. countries with mild winters, but it can endure light
frost without much permanent damage. Its attrac-
tive, soft foliage makes it a desirable small tree and it
Distribution
should be cultivated more often.
Australia: New South Wales, South Australia, Victo-
ria, Western Australia.
TDWG codes: 50 NSW-NS SOA VIC WAU-WA Callitris rhomboidea R. Br. ex Rich. & A. Rich.,
in A. Richard (ed.) Comm. Bot. Conif. Cycad.: 47.
1826. Type: Australia: New South Wales, Sydney,
Ecology
Port Jackson, R. Brown 3107 (holotype BM). Fig. 68
In coastal and inland dunes, as well as on sandy
floodplains, hillsides, low ridges (‘breakaways’) and
Etymology
laterite (hardpan). In coastal dune areas it may form
more or less dense stands excluding other trees and The species epithet refers to the rhomboid shape of
shrubs, or it associates with Acacia sp. and Mela­leuca the seed cone scales.
lanceolata; elsewhere it occurs in low Eucalyptus
woodland (‘mallee’), associated with other Myrta­
Vernacular names
ceae (e.g. Melaleuca, Leptospermum), Proteaceae (e.g.
Banksia, Dryandra, Grevillea, Hakea), Acacia spp., Port Jackson Pine, Oyster Bay Pine, Cypress-pine,
Allocasuarina sp., Dodonaea sp., and Callitris spp.; Illawarra Mountain Pine
in (dwarf) scrubland (‘kwongan’ in SW Australia)
with Myrtaceae (‘Melaleuca thicket’), Proteaceae,
Description
and Xanthorrhoea thorntonii; in inland dunes and
dune slacks with Eucalyptus spp., Acacia spp., Myrta­ Shrubs or trees to 10–15 m tall; multistemmed or
ceae, and Callitris verrucosa; on sandstone hillsides monopodial, usually branching low and often fas-
and ridges with Eucalyptus spp., Acacia spp., Allo- tigiate, up to 40 cm d.b.h. Bark soon scaly, on the
casuarina cristata, Dodonaea sp., Callitris columel- lower trunk shallowly fissured, hard, slowly ex­
laris, C. rhomboidea, and C. ­endlicheri. The altitude foliating in small flakes, dark grey-brown, inner lay-
ranges from near sea level to 670 m a.s.l. Soils vary ers brown. Branches spreading or ascending at ca.
from calcareous, humus-rich old coastal or ripar- 45°, or fastigi­ate, long and slender, forming a conical
ian sand dunes, (where it develops to a monopodial or (broad) pyramidal, dense or more or less open
or multi-stemmed tree) to white, yellow, brown or crown. Foliage branchlets spreading or ascending
red sand or loam flats over limestone, sandstone or to erect, ultimately very slender, triangular in cross
laterite (hardpan; on this it develops only to a me- section, 0.6–1 mm diam., covered with closely ap-
dium height shrub); also in disturbed road verges pressed leaves, persistent. Leaves in alternate whorls
through arable land or pasture. The climate is mild of 3, decurrent, closely appressed, apices appressed
to hot and characterized by dry periods (which can or free especially on some leading shoots (whip
 shoots), connate but parting with thickening of ­ anthorrhoea sp., Restio sp., Pteridium sp., ­Callitris
X
branches, persisting several years, linear, 3–6 mm ­endlicheri, and C. preissii. Soils are (‘skeletal’) white
long on ultimate branchlets, 0.3–0.7 mm wide, abax- or grey sand over sandstone, limestone, shale, gran-
ially keeled; margins denticulate near slightly broad- ite (Adamellite), or it is growing in talus, rocky creek
ened acute-acuminate apex; epistomatic, stomata in bottoms, or crevices of bedrock; also in fields (sheep
two narrow, marginal bands; abaxial surface weakly pasture), with invasives (e.g. Ulex europaeus in Tas-
verrucose, yellowish green to glaucous green. Pol- mania) as indicators of disturbance. The altitudinal
len cones solitary or paired and terminal on ulti- range is from near sea level to ca. 1250 m a.s.l. The
mate branchlets, ovoid-globose to oblong, 1.5–2.5 climate is mild to temperate, mesic or with summer
× 1–2 mm, yellowish green turning orange-brown; rainfall in the north and winter rainfall in the south
240 microsporophylls 8–14, in alternate whorls of 3, pel- of its range; frost and snow can occur at higher
tate, with denticulate margins, bearing 3–4 abaxial ­altitudes.
pollen sacs near the lower margin. Seed cones ter-
minal on short, thick, leafy shoots, solitary or more
Conservation
commonly aggregated in dense, compact clusters
on branches, maturing in 1–1.5 years, persistent, IUCN: LC
depressed-­globose when closed, 12–15 × 13–17 mm,
angular with large, protruding umbos, smooth, of-
Uses
ten lustrous, becoming finely rugose when open,
blackish brown or nearly black, weathering grey. As a widespread species its timber, being relatively
Bract-scale complexes in 2 alternate whorls of 3, small, was mainly used for fence posts until metal
only partly incised towards cone base when opened; replaced wood for this pupose in farming. It has
upper ones largest, 10–13 × 8–12 mm, tapering to been introduced in New Zealand, where it has be-
an obtuse apex; smaller ones narrower 6–8 × 5–7 come naturalized near Auckland. This introduction
mm; forming a prominent and acute umbo below had no horticultural purpose as far as I know. The
the apex of each scale; adaxial surface dark purplish species remains relatively rare in botanic gardens
brown with light seed marks. Columella trilobed to and arboreta, although it is available in the UK nurs-
tripartite, 3–8 mm wide, obtuse. Seeds 4–10 on each ery trade (Grimshaw & Bayton, 2009: 186).
scale, ovoid or triangular, flattened, up to 8 mm in-
cluding wings (seed body ca. 3 × 2 mm), dark brown
to black with large, greyish concave hilum; wings Callitris roei (Endl.) F. Muell., Syst. Census Austral.
2–3 on sides, 2–3 mm wide, more or less equal in Pl. 1: 109. 1882. Frenela roei Endl., Syn. Conif.: 36.
shape and size. 1847. Type: Australia: Western Australia, along the
road from Newdegate to Lake King, 35 km E of
Newdegate, A. Farjon 375 (neotype K). Fig. 69, 70
Distribution
Australia: New South Wales, Queensland, South
Etymology
Australia, Tasmania, Victoria.
TDWG codes: 50 NSW-NS QLD-QU SOA TAS VIC This species was named after J. S. Roe (1797–1878),
who collected the original specimens, later de-
stroyed at W.
Ecology
Commonly in open woodland or scrubland on
Vernacular names
tablelands or mountains, locally on riparian or
coastal sands, often amongst boulders; associated Roe’s Cypress-pine
with Eucalyptus spp., other Myrtaceae (e.g. Kunzea,
Lepto­spermum), Proteaceae (e.g. Banksia, Cono-
Description
spermum, Hakea), Acacia spp., Allocasuarina spp.,
Daviesia ­ulicina, Dilwenia sp., Leucopogon spp., Shrubs or small trees 1–5 m; multistemmed or mo-
Prostanthera lasianthos, Spyridium sp., Hibbertia sp., nopodial, branching low, up to 20 cm d.b.h. Bark on
the lower trunk thin, shallowly fissured, hard, slowly
Ecology
exfoliating in small flakes, grey. Branches spreading
or ascending at ca. 45°, long and pliant, forming a In low Eucalyptus woodland (‘mallee’) with Eucalyp-
broad, rounded, dense or more or less open crown. tus spp., other Myrtaceae (e.g. Melaleuca), Proteace-
Foliage branchlets rigid and thick, ultimately twist- ae (e.g. Banksia, Dryandra, Hakea), Leschenaultia
ed and of unequal length, triangular in cross section, sp., Callitris preissii, and occasionally in open dwarf
1.2–1.5 mm diam., covered with closely appressed scrubland (‘kwongan’); on (gravelly) white or grey
leaves, persistent. Leaves in alternate whorls of 3, sand, loam or clay and on laterite (hardpan), also in
decurrent, closely appressed, apices appressed or disturbed road verges. The altitudinal range is from
some free especially on some leading shoots (whip 50 m to 350 m a.s.l. The climate is characterized by
shoots), connate but parting with thickening of warm, dry summers and winter rainfall. 241
branches, linear, 2–5 mm long on ultimate branch-
lets, 1–1.5 mm wide, abaxially prominently keeled;
Conservation
margins denticulate near slightly broadened obtuse
or acute apex; epistomatic, stomata in two narrow Due to widespread conversion of the original veg-
marginal bands, a few stomata visible near the apex; etation cover to arable farming this species has suf-
abaxial surface weakly verrucose, yellowish green fered a substantial reduction in its area of occupancy
to light green. Pollen cones solitary and terminal in the 20th century. Remaining large reserves are
on ultimate branchlets, oblong to cylindrical, 3–6 mainly concentrated along the south coast, while
× 1.2–2 mm, yellowish green turning light brown; in the interior these are small and scattered, or have
microsporophylls 10–16, in alternate whorls of 3, a limited protective status as road verge ‘reserves’
peltate, broadly rhombic, with erose-denticulate of semi-natural vegetation. Habitat degradation is
margins, bearing 3–4 abaxial pollen sacs near the mainly caused by increasing salinity of soils in the
lower margin. Seed cones terminal on short, thick, region; along roads fires also have increased, but it is
leafy shoots, mostly aggregated in dense, compact unknown how this species reacts to this. It appears
clusters on stems and branches, maturing in 1–1.5 in some situations to react positively to mechanical
years, persistent, depressed-globose when closed, soil disturbance along roads (personal observations,
(10–)12–18 × 12–20 mm, with concave lower parts 1997) and should be at least mildly tolerant of fire.
of scales, smooth, finely rugose when open, whitish IUCN: VU (A1c)
grey to metallic grey, weathering dull grey. Bract-
scale complexes in 2 alternate whorls of 3, open-
Uses
ing not very wide; upper ones largest, (10–)12–15
× 6–10 mm, with parallel sides and obtuse apex; There are no economic uses recorded of this species.
smaller ones wider at base, triangular, with acute
apex; a prominent and acute umbo below the apex
of each scale; abaxial surface concave, becoming ru- Callitris sulcata (Parl.) Schltr., Bot. Jahrb. Syst. 39:
gose, grey; adaxial surface yellowish or more often 16. 1906. Frenela sulcata Parl., Index Sem. Hort.
red-brown with light seed marks. Columella large, Florent. 1862: 23. 1862; Nothocallitris sulcata (Parl.)
triangular or trilobed, 3–6 mm long and wide, ob- A. V. Bobrov & Melikyan, Komarovia 4: 85. 2006.
tuse. Seeds 3–6 on each scale, ovoid or triangular, Type: New Caledonia: Grande Terre, Province Sud,
flattened, up to 10 mm including wings (seed body C. Moore 5 (holotype K).
3–5 mm long), yellowish or red-brown, with large,
whitish concave hilum; wings 2 on each side, up to
Etymology
8 × 4 mm, unequal in shape and size.
The species epithet (Latin sulcatus = ‘grooved’) re-
fers to the appearance of the branchlets.
Distribution
SW Western Australia, near the south coast and in
Vernacular names
parts of the Wheat Belt region.
TDWG codes: 50 WAU-WA Sapin de Comboui (French)
 Description Taxonomic notes
Shrubs or small trees to 6–10(–12) m tall; trunk mo- The two species of Callitris in New Caledonia are
nopodial in trees, sometimes forked, branching low, probably relicts of the Australian connection about
up to 40 cm d.b.h. Bark soon scaly, to 10–20 mm 80 Ma, and somewhat resemble C. macleayana in
thick, fissured, stringy, exfoliating in long strips, foliage characters. They retain a 4-whorled phyllo-
light brown weathering grey. Branches spread- taxis with juvenile acicular leaves into semi-mature
ing or ascending, forming an irregular, broad and or mature plants. In C. sulcata the transition be-
usually open crown. Foliage branchlets in tufts at tween leaf types is abrubt, but phyllotaxis changes
ends of main branches, assurgent or erect, ultimate from 4 to 3 whorls earlier than leaf type. Cones are
242 branchlets long, apprearing conspicuously articulate only borne on branchlets with 3-whorled phyllotaxis
with scale leaves, triangular in cross section, ca. 1.5 and mature type leaves.
mm diam., covered with closely appressed leaves,
green, persistent. Leaves of two types on plants up
Distribution
to 1 m tall. Adult (scale) leaves in alternate whorls
of 3, decurrent, closely appressed, apices appressed New Caledonia, Grande Terre, Province Sud, in val-
or some free especially on some leading shoots leys of the Comboui, Dumbea and Tontouta Rivers
(whip shoots), connate but parting with thicken- and some tributaries.
ing of branches, linear, 3–7 mm long on ultimate TDWG codes: 60 NWC
branchlets, 0.7–1 mm wide, obtusely keeled abaxi-
ally; margins erose-denticulate near obtuse or acute
Ecology
apex; epistomatic, stomata in two narrow marginal
bands; abaxial surface smooth, green. Pollen cones On serpentine on slopes or in valleys close to rivers
solitary and terminal on ultimate branchlets, ovoid and creeks; in scrubland on ultramafic soils (‘maquis
or ovoid-oblong, 3–5 × 1.5–2.5 mm, yellowish green minier’) and in the ecotone towards low rainforest
turning light brown; microsporophylls 9–15, in al- (gallery forest), at altitudes between 40 m and 200
ternate whorls of 3, peltate-triangular, with erose- m a.s.l. It grows in thickets, or scattered with other
denticulate margins, obtuse or acute, bearing 4–6 conifers e.g. Dacrydium araucarioides, D. balansae,
abaxial pollen sacs near the lower margin. Seed Neocallitropsis pancheri and Podocarpus novae-
cones terminal on short, thin, leafy shoots, solitary, caledoniae and numerous ferns and angiosperms.
maturing in 1 year, caducous (or persistent a short The climate is tropical, with high levels of rainfall
time after seed dispersal), ovoid-globose to subglo- throughout the year.
bose when closed, 8–11 × 7–10 mm, with concave
lower parts of scales when open, smooth, finely ru-
Conservation
gose when open, greenish maturing to dull brown.
Bract-scale complexes in 2 alternate whorls of 3, not This species is limited to a few river valleys in S New
opening very wide; upper ones only slightly larger, Caledonia and their tributaries and headwaters,
7–10 × 5–6 mm, oblong, thick; lower ones 4–5 mm where it is rare. Timber extraction is reported from
wide; a prominent and acute umbo below the apex the Comboui Valley, and although this species is not
of each scale; adaxial surface rugose, red-brown or an important timber tree, it is likely to be felled with
dark brown with inconspicuous seed marks. Colu- other trees. It is vulnerable to fire and disturbance
mella relatively large, 3–4 mm long, irregularly both of which have increased in recent years. Regen-
angular, 1.5–2 mm wide, narrowed at base. Seeds eration appears to be poor in several localities. Pro-
usually 2 at base of each upper scale, ovoid or trian- tection in ad hoc river bank reserves is inadequate
gular, flattened, 4–5 × 3 mm, light brown, with very and ex situ propagation is required to back up in situ
small hilum; wings 2 on each side, up to 7 × 2.5 mm, regeneration in disturbed sites.
unequal in shape and size. IUCN: EN [B1ab(iii)+2ab(iii), C2a(i)]
 green turning light brown; microsporophylls 10–16,
Uses
decussate or some in alternate whorls of 3, peltate,
No uses of any kind have been reported of this spe- orbicular, with erose-denticulate margins, bearing
cies. 2–3 abaxial pollen sacs near the lower margin. Seed
cones terminal on short, thick, leafy shoots, solitary
or more often clustered on branches and stems, ma-
Callitris verrucosa (A. Cunn. ex Endl.) F. Muell., turing in 1–1.5 years, persistent, subglobose to ovoid-
Rep. Burdekin Exped.: 19. 1860. Frenela verrucosa globose when closed, 18–30 × 15–25 mm, nearly
A. Cunn. ex Endl., Syn. Conif.: 37. 1847; Callitris smooth when immature, becoming densely covered
preissii Miq. subsp. verrucosa (A. Cunn. ex Endl.) with small (1.5–2.5 mm) verrucae, purplish grey to
J. Garden, Contr. New South Wales Natl. Herb. 2 blackish, weathering grey. Bract-scale complexes in 243
(5): 375. 1957; Callitris preissii Miq. var. verrucosa 2 alternate whorls of 3, deeply incised towards the
(A. Cunn. ex Endl.) Silba, Phytologia Mem. 7: 17. cone base when opened, thick; upper ones largest,
1984. Type: Australia: New South Wales, Lachlan 15–25 × 7–12 mm, acute or obtuse; smaller ones nar-
River, A. Cunningham 372 (holotype K). Pl. 9 rower; bracts entirely included, rarely showing the
bract tip below the scale apex; adaxial surfaces red-
dish brown or dark brown with light seed marks.
Etymology
Columella variable, conical, or bi- to tripartite, 2–3
The species epithet refers to the outer surface of the × 1.5–2 mm, acute. Seeds 6–12(–16) on each scale,
seed cone scales, which is covered with verrucae or triangular to ovoid, flattened, up to 9 mm including
small blisters. wings (seed body up to 3–5 × 2–3 mm), dark brown
or red-brown with whitish concave hilum; wings
2(–3) on opposite sides, 2–4 mm wide, more or less
Vernacular names
equal or unequal in shape and size.
Mallee Pine, Cypress Pine, Turpentine Pine, Scrub
Cypress-pine, Camphor Wood
Taxonomic notes
The species rank of this taxon has not been accepted
Description
by several authors (Garden, 1957; Harden & Thomp-
Shrubs or trees to 8–10(–15) m tall; multistemmed, son, 1990). Maiden (1907) included C. tuberculata in
or monopodial but then usually forked, up to 0.6 m this species, while Baker & Smith (1910) treated the
d.b.h. Bark soon scaly, on large trunks thick, deep- two as separate species, as has been done recently
ly fissured, stringy, exfoliating in large strips, light in Flora of Australia 48 (1998). The classification of
brown; outer bark grey. Branches spreading or as- C. verrucosa as an infraspecific taxon of C. preissii
cending, long, forming a sympodial, broadly domed, (Garden, 1957) is, in my view, based on a misinterpre-
more or less open and irregular crown, or a rounded tation of morphological variability. While C. preissii
shrub. Foliage branchlets ascending or erect, ulti- is a species with high polymorphism, to some ex-
mately very slender, terete, 0.6–0.8 mm diam., cov- tend correlated with environmental factors, C. ver-
ered with closely appressed leaves, persistent. Leaves rucosa is very constant in most of its morphological
in alternate whorls of 3, decurrent, closely appressed character states and traits throughout its range. It
including apices (except on some whip shoots), con- does not occur anywhere on the coast and its trees/
nate but parting with thickening of branches, linear, shrubs on dunes are not dissimilar from those on
1.5–3 mm long on ultimate branchlets, 0.5–1 mm level hardpan plains. While growth on dunes is often
wide, abaxially convex; margins denticulate near better than on hardpan, C. verrucosa never devel-
obtuse or acute apex; mostly epistomatic, abaxial ops into a monopodial tree in either environment.
stomata near base of leaves only or usually absent, The single true variation I have observed in the field
adaxial stomata in two marginal bands and a few is the occurrence of two foliage colour phases, yel-
below the apex; abaxial surface smooth, yellow- lowish green and glaucous green, growing even side
ish green or glaucous green. Pollen cones solitary by side in some populations in Wyperfeld National
or more often 2–3 together, terminal on ultimate Park (field observations A. Farjon & A. Watt, 1997).
branchlets, cylindrical, 2–3.5 × 1–1.5 mm, yellowish The verrucae on the mature cones (growing cones
 8

244

1
5

2
3

Pl ate 9 . Callitris verrucosa. 1. Branch with foliage and seed cone. 2. Section of branchlet with leaves.
3. Branchlet with pollen cones. 4. Microsporophyll with pollen sacs. 5. Immature seed cone. 6. Branch with
seed cones. 7, 8. Seeds.
are nearly smooth) are densely set, small and of near perate to subtropical and characterized by long dry
constant size in all live plants and herbarium speci- periods and short periods of rainfall.
mens seen. These characters are good indicators of
a good biological species that is partially sympatric
Conservation
with C. preissii, as here circumscribed, but remains
separate. Some species are morphologically (and ge- This widespread species occurs mostly in semi-
netically) much less variable than others, even when deserts and inland dune areas, where it is currently
they have an extensive range like this species. not threatened with extinction. In certain regions,
e.g. on the Murray River plain, irrigation agriculture
may have destroyed populations replacing the origi-
Distribution
nal vegetation with arable or viticulture. Clearance 245
Australia: New South Wales, Queensland (Darling for wheat growing in Victoria and South Australia
Downs), South Australia, Victoria, interior of West- will have destroyed other populations. It is not pos-
ern Australia. sible with the available data to determine if the area
TDWG codes: 50 NSW-NS QLD-QU SOA VIC WAU- of occupancy of this species has been significantly
WA reduced, but its extensive range and inclusion in
large protected and/or unused areas provides an as-
surance of continued survival.
Ecology
IUCN: LC
In low Eucalyptus woodland (‘mallee’) or in (open)
scrubland, associated with Eucalyptus spp., other
Uses
Myrtaceae (e.g. Baeckea, Leptospermum, Melaleuca),
Proteaceae (e.g. Banksia, Grevillea, Hakea), Acacia The wood of this species has similar properties to
spp., Allocasuarina sp., Beyeria opaca, Leucopogon that of its congeners, but the shrubby habit makes it
sp., Eriostemon sp., Dillwynia sp., Hibbertia sp., Spy- unsuitable for timber. It is not known in cultivation
ridium sp., Lomandra juncea, ‘spinifex’ hummocks outside a few botanic gardens. There are two foliage
(Triodia sp.), and Callitris preissii; on white, yellow, colour forms occurring in the wild, often together:
brown or red sand dunes and flats, sometimes on yellowish green and glaucous green; these could be
sand over limestone or granite. Its altitudinal range selected and enhanced under cultivation.
is from 30 m to 520 m a.s.l. The climate is warm tem-
 Calocedrus Kurz, J. Bot. 11: 196. 1873. Type: Calocedrus macrolepis Kurz
(Cupressaceae).

Greek: calo = beautiful, so ‘beautiful cedar’; cedrus 2a. Seed cones on long branchlets with numerous
is a classical name applied to certain conifers with modified monomorphic scale leaves, 10–22 ×
fragrant wood, it is not clear which! 4–7 mm; normally 1 seed developing on each
fertile scale C. macrolepis
2b. Seed cones on short branchlets with few modi-
Description
fied scale leaves, less than 16 mm long; 1 or 2
246 Evergreen, monoecious trees with a monopodial seeds developing on each fertile scale 3
trunk. Resin cavities in leaves. Bark fissured or fur- 3a. Leaves and seed cones very small, the latter
rowed, fibrous, stringy, exfoliating in long plates or (4–)5–6(–7) × (2.5–)3–4 mm; normally 2 seeds
strips. Branches curved down or spreading to as- developing on each fertile scale C. rupestris
surgent, forming a pyramidal or sympodial crown. 3b. Leaves and seed cones larger, the latter 10–15 ×
Fastigiate forms mostly restricted to trees in culti- 4–6 mm; normally 1 seed developing on each
vation. Foliage branches plagiotropic; (sub)ultimate fertile scale C. formosana
branchlets flattened, covered with scale leaves; later-
al branchlets continually deciduous. Leaves decus-
sate, appearing in whorls of 4, decurrent, imbricate, Calocedrus decurrens (Torr.) Florin, Taxon 5 (8):
dimorphic, with oblong to obtrullate facials and 192. 1956. Libocedrus decurrens Torr., Smithsonian
slightly broader, conduplicate laterals, very variable Contr. Knowl. 6 (2): 7. t. 3. 1853. Type: USA:
in size dependent on growth of branchlets; margins California, Shasta Co., [“Upper waters of the
entire; apices incurved or recurved, those of lateral Sacramento River”], J. C. Frémont 492 (holotype
leaves meeting those of facials; stomata mostly on US?, isotype MO). Fig. 71, 72
the underside of lateral leaves in two lines or bands,
glands inconspicuous near apex of facials, absent
Etymology
on laterals. Pollen cones solitary, terminal, cylindri-
cal; microsporophylls (6–)8–14, decussate, with 3–5 The species epithet refers to the decurrent leaves on
abaxial pollen sacs. Seed cones solitary, terminal, primary shoots.
10–35 × 4–13 mm, caducous. Bract-scale complexes
in 3(–4) decussate pairs, the proximal pair small, re-
Vernacular names
curved, sterile, the middle pair spreading, fertile, the
distal pair fused, sterile. Seeds 1–2 per fertile scale, Incense Cedar, California Incense Cedar, White Ce-
with 2 very unequal wings. Seedlings with 2 coty- dar, Californian Post Cedar, Bastard Cedar; Cedro
ledons. incienso (Spanish)

4 species.
Description
Trees to 60(–69) m tall; trunk monopodial, broadly
Distribution
based or buttressed, up to 3–4.5 m d.b.h. Bark on
W coast of North America, from Oregon to Baja lower trunk deeply furrowed, fibrous, stringy, ex-
California; Taiwan, SW China, mainland SE Asia. foliating into long plates or strips, dark red-brown.
Branches curved down below, then spreading, as-
surgent near the top, long persistent; higher order
Key to the species of Calocedrus
branches pendulous below, forming a columnar to
1a. Seed cones when mature 20–35 mm long; nor- pyramidal, eventually domed and more open crown.
mally 2 seeds developing on each fertile cale Foliage branches numerous, plagiotropic, spreading
C. decurrens or erect near the top, ultimately flattened, covered
1b. Seed cones when mature (4–)5–22 mm long; 1 with green leaves, lateral branchlets deciduous in
or 2 seeds developing on each fertile scale 2 2–3 years. Leaves decussate, appearing in whorls
of 4, imbricate; laterals partly covering facials, de- locally with Sequoiadendron giganteum, Chamae-
current, (weakly) dimorphic, conduplicate, linear- cyparis lawsoniana, Tsuga heterophylla or Thuja
lanceolate, with slightly incurved acute apex; facials plicata, and in drier southern sites with Pinus coul-
slightly shorter than laterals, oblong to obtrullate, teri and Pseudotsuga macrocarpa. The undergrowth
obtuse-acuminate; leaves persisting several years, of these mixed conifer forests varies mostly with
variable in size, from 2 × 1.5 mm on ultimate lateral altitude and edaphic conditions and is diverse, es-
branchlets to 15 × 3 mm on leading (whip) shoots; pecially on ultramafic rocks, with Arctostaphylos
margins entire; amphistomatic, abaxial stomata patula, A. viscida, Ceanothus cordulatus, C. inte-
on laterals in two narrow bands, more numerous gerrimus, C. parvifolius, Castanopsis sempervirens,
on the underside, stomata on facials restricted to Gaultheria shallon and many other shrubby species.
sides and hidden in grooves between leaves; glands In most conifer forest associations C. decurrens is a 247
weakly developed, most conspicuous near apex of relatively minor component, where it often occupies
facials, all exposed faces light green or sometimes open canopy stands on hot, dry sites. In the Sierra
dark green. Pollen cones solitary and terminal on Nevada Mixed Conifer Forest it may play a much
more or less pendulous branchlets, oblong, 6–8 × greater role in the canopy locally. Other forest types
2–3 mm, yellow turning brown; microsporophylls include also Quercus spp., Castanopsis chrysophylla,
10–14, ­decussate, peltate-orbicular with denticulate Lithocarpus densiflorus and Arbutus menziesii, to-
margins and acutish apex, bearing 3–4 abaxial pol- gether with conifers. The altitudinal range of Calo-
len sacs near lower margin. Seed cones terminal cedrus decurrens is from 50 m to 2010 m a.s.l. in the
on more or less pendulous branchlets with mostly north and between 910 m and 2960 m a.s.l. in the
unmodified leaves (except for 3–4 subtending pairs south of its range. This species is rather tolerant to
of small scale leaves), solitary, often numerous, ma- soil types, with a huge range of pH values, but the
turing within 1 year, caducous, ovoid-oblong or ob- soil usually well drained; it is only rare on limestone.
long when closed, (15–)20–35 × 8–13 mm, smooth or It tolerates hot, dry summers well, but is equally in-
striated, turning red-brown. Bract-scale complexes sensitive to frost and snow cover.
in 3(–4) decussate pairs; proximal pair undevel-
oped and sterile, with recurved bracts; middle pair
Conservation
spreading wide, fertile, slightly curved, abaxially
convex, with a minute subapical bract tip and 2 de- IUCN: LC
pressed seed marks near the base of each scale; distal
pair infertile, straight, laterally flattened and fused to
Uses
a flat plate with 2 small seed marks and apical bract
tips; latter two pairs 18–30 × 7–10 mm, their adaxial Incense cedar is an important timber tree. The wood
faces rugose, light brown. Seeds oblong, slightly flat- is used for the manufacture of pencils and in build-
tened, tapering to apex, 8–12 × 3–4 mm, light whit- ing for exterior siding and windows of houses and
ish brown, with 2 wings of unequal shape and size; garden amenities like fences and trellises. As with
largest wing 18–23 mm longwing; smallest a narrow many taxa in this family, the wood is decay resistant,
strip with a free apex. which makes it especially useful for these purposes
in the wetter coastal areas of the Pacific States in
the USA. It is also widely planted as an ornamental
Distribution
tree and only then often grows into a fastigiate habit
USA: California, Oregon; NW Mexico: Baja Califor- (probably mostly as the cultivar ‘Columnaris’ which
nia Norte. was named by Beissner Libocedrus decurrens var.
TDWG codes: 73 ORE 76 CAL 79 MXN-BC ­columnaris but does not occur in nature). Incense
cedar performs well in urban settings as it is relative-
ly tolerant of air pollution. In horticulture a number
Ecology
of cultivars are known, especially with variegated
In mixed conifer forest with Pinus jeffreyi, P. pon- or differently coloured foliage. Despite its common
derosa, P. lambertiana, P. monticola, Abies concolor, name, it is not used for incense burning, although its
A. grandis, A. magnifica, and Pseudotsuga menziesii, foliage is fragrant.
 Calocedrus formosana (Florin) Florin, Taxon 5 (8): minal on more or less pendulous branchlets with
192. 1956. Libocedrus formosana Florin, Svensk Bot. ultimately slightly modified (shortened) leaves, soli-
Tidskr. 24: 126. 1930; Calocedrus macrolepis Kurz tary, often numerous, maturing within 1 year, cadu-
var. formosana (Florin) W. C. Cheng & L. K. Fu, cous, oblong when closed, 10–15 × 4–6 mm, smooth
Fl. Reipubl. Pop. Sin. 7: 327. 1978. Type: Taiwan: or slightly rugose, often glaucous, turning brown.
[“Taihoku province” (planted)], E. H. Wilson 10960 Bract-scale complexes in 3 decussate pairs; proxi-
(lectotype K). Fig. 73 mal pair undeveloped and sterile, with recurved
bracts; middle pair spreading slightly, fertile, slight-
ly curved, abaxially convex, with a small subapical
Etymology
recurved bract tip and 1 depressed seed mark near
248 The species epithet indicates Taiwan, formerly base; distal pair infertile, straight, laterally flattened
known as Formosa; see also Latin: formosus = hand- and fused to a flat plate with apical bract tips; latter
some or well formed. two pairs 10–13 × 4–5 mm, their adaxial faces soft,
smooth, light brown. Seeds 1(–2) per cone, ovoid,
flattened, acutish at apex, 4–5 × 1.8–2 mm, brown,
Vernacular names
with 2 wings of unequal shape and size; largest wing
Taiwan Incense-cedar; Taiwan cui bai (Chinese) 10–12 mm long; smallest wing a narrow strip with a
free apex or virtually absent.
Description
Taxonomic notes
Trees to 20–25 m tall; monopodial, up to 2–3 m
d.b.h. Bark on lower trunk furrowed, fibrous, ex- This taxon was treated as a variety of C. macrolepis
foliating into long plates or strips, red-brown. in Flora of China 4 (1999); in the Flora of Taiwan (Li,
Branches often few and thick, spreading, ascending 1975) C. formosana is recognized as a species. The
near the top, long persistent, higher order branch- morphological differences are small but constant
es spreading, forming a pyramidal, eventually and C. formosana only occurs in Taiwan.
broad and more or less flat-topped crown. Foliage
branches numerous, plagiotropic, branching alter-
Distribution
nately, ultimately flattened and articulate, covered
with green leaves, lateral branchlets deciduous in Taiwan: central and northern mountains.
2–3 years. Leaves decussate, appearing in whorls TDWG codes: 38 TAI
of 4, imbricate, dimorphic; laterals partly covering
facials, decurrent, conduplicate, linear-lanceolate,
Ecology
with recurved-incurved acute apex; facials oblong
to obtrullate, obtuse-acuminate; leaves persisting This species occurs in mixed conifer-broad-leaved
several years; variable in size, 1–5 mm long on (sub) climax forest, often as an emergent, associated with
ultimate lateral branchlets to 15 × 3 mm on leading Pseudotsuga sinensis, Taiwania cryptomerioides,
(whip) shoots; margins entire; amphistomatic, abax- Castanopsis spp., and Cyclobalanus (= Quercus) spp.
ial stomata on laterals in two lines or bands, very few in deep, rich forest soil over shalestone or schist; also
or absent on upperside, much more numerous on in secondary forest and on cliffs and rocky ridges. Its
underside, stomata on upperside facials restricted to altitudinal range is (300–)800–2000 m a.s.l.
hidden margins, on underside facials in two bands;
glands inconspicuous near apex of facials or absent;
Conservation
exposed faces green or sometimes dark green, with
glaucous green stomatal bands. Pollen cones solitary This species extends in an area of less than 5000 km2
and terminal on ultimate branchlets, oblong, 4–5 × and is limited to under five isolated locations while it
2–2.5 mm, reddish turning brown; microsporophylls suffers a continuing decline in its area of occupancy.
(6–)8–10, decussate, peltate-orbicular with denticu- The causes are timber cutting, transformation of old
late margins and acutish apex, bearing 2–3 abaxial growth forest to managed production forest, and ex-
pollen sacs near the lower margin. Seed cones ter- pansion of agriculture, especially livestock ­grazing.
Some stands are now protected in reserves and there 15 × 3 mm on leading (whip) shoots; margins en-
is a programme of ex situ conservation to back up tire; amphistomatic, abaxial stomata on laterals in
the remaining populations. two lines or bands, very few or absent on upperside,
IUCN: EN (B1+2b) much more numerous on underside, stomata on
upperside facials restricted to hidden margins, on
underside facials in two bands; glands inconspicu-
Uses
ous near apex of facials or absent, exposed faces
The valuable wood of this tree is in demand for con- light green with whitish green stomatal bands. Pol-
struction purposes, mainly at a local or regional lev- len cones solitary and terminal on ultimate branch-
el. It is rare in cultivation despite the fact that it can lets, oblong, 4–8 × 2–3 mm, purplish red turning
survive winters in e.g. England provided it is planted light brown; microsporophylls (8–)10–14, decus- 249
in a sheltered spot; a few recently planted trees at the sate, peltate-orbicular with denticulate margins and
Royal Botanic Gardens, Kew have seed cones and obtuse-acuminate apex, bearing (3–)4(–5) abaxial
are doing well. Although known botanically since pollen sacs near the lower margin. Seed cones ter-
1930, it was apparently introduced to cultivation minal on pendulous (5–)10–20 mm long branchlets
only recently (Grimshaw & Bayton, 2009: 187). with modified (shortened) monomorphic leaves in
decussate pairs, solitary, often numerous, matur-
ing within 1 year, caducous, oblong when closed,
Calocedrus macrolepis Kurz, J. Bot. 11: 196., t. 133. 10–22 × 4–7 mm, smooth or slightly rugose, often
1873. Libocedrus macrolepis (Kurz) Benth. & Hook. glaucous, turning brown. Bract-scale complexes
f., Gen. Pl. 3 (1): 426. 1880. Type: China: Yunnan, in 3(–4) decussate pairs; proximal pair undevel-
Daying River, Hotha, D. J. Anderson s.n. (holotype oped and sterile, with recurved bracts; middle pair
K). Fig. 74 spreading slightly, fertile, nearly straight, abaxially
convex, with a small subapical recurved bract tip
and 1(–2) depressed seed marks near the base; distal
Etymology
pair infertile, straight, laterally flattened and fused
The species epithet (Greek/Latin: macro = large; to a flat plate with apical, curved bract tips; the lat-
lepis = scale) refers to the size of the seed cone scales. ter two pairs 10–20 × 4–6 mm, adaxial faces soft,
smooth, light brown. Seeds 2(–4) per cone, ovoid-
oblong, slightly flattened, acute at apex, 5–6 × 2–3
Vernacular names
mm, brown, with 2 wings of unequal shape and size;
Chinese Incense-cedar; cui bai (Chinese) largest wing 12–15(–20) mm long; smallest wing a
narrow strip with a free apex or virtually absent.
Description
Distribution
Trees to 30–35 m tall; monopodial, up to 1.5–2 m
d.b.h. Bark on lower trunk fissured, exfoliating into China: Guangxi (Jingxi Xian), Guizhou, Guang-
long plates or strips, pale grey-brown. Branches of- dong?, Hainan Island, Yunnan; Myanmar [Burma],
ten few and thick, spreading, ascending near the Thailand, Laos, Vietnam.
top, long persistent, higher order branches spread- TDWG codes: 36 CHC-GZ CHC-YN CHH CHS-GX 41
ing, forming a pyramidal, eventually sympodial, LAO MYA THA VIE
broad, rounded or flat-topped crown. Foliage
branches numerous, plagiotropic, branching alter-
Ecology
nately, ultimately flattened and articulate, covered
with green leaves, lateral branchlets deciduous in Relatively common in montane mixed evergreen
2–3 years. Leaves decussate, appearing in whorls conifer-broad-leaved forest dominated by Fagaceae
of 4, imbricate, dimorphic; laterals partly covering and with scattered conifers, e.g. Cunninghamia,
facials, decurrent, conduplicate, linear-lanceolate, Keteleeria and Pinus; also in rocky places and much
with recurved-incurved acute apex; facials oblong planted in roadsides and field margins. The altitudi-
to obtrullate, obtuse-acuminate; leaf size variable, nal range is from ca. 1000 m to 2000 m a.s.l.
1.5–8 mm long on (sub)ultimate lateral branchlets to
 whorls of 4, decurrent at base, broadly obtuse to
Conservation
obtuse at apex, dimorphic; facial pair flattened, (1–
This species is under threat from cutting for both )2–6(–7) mm long, (1.5–)2–2.5 mm wide; lateral pair
timber and firewood, especially in more accessible conduplicate, boat-shaped, (1.5–)2–6(–7) mm long,
areas in Yunnan. In that province it is still fairly (0.3–)0.5–0.75(–1) mm wide, their margins over-
common; in its outlying stations, e.g. Hainan Island lapping with facial leaves, eglandular. Pollen cones
and Vietnam, it is more scarce to rare. Protective terminal, solitary, cylindrical, (4.5–)5–6 mm long,
measures were suggested in the form of reserves as 1.5–2(–2.2) mm wide; microsporophylls decussate,
well as ex situ cultivation in Fu & Jin (eds., 1992). in 8–11 pairs (the lowest 2–4 pairs sterile), 0.8–1.2
IUCN: VU (B1+2b) mm long, 1–1.2 mm wide, with minutely erose mar-
250 gins, broadly obtuse or rounded at apex, light green
turning light brown, each with 2–6 broadly ovoid
Uses
to subglobose, 0.3–0.4 mm wide pollen sacs. Seed
In China this species is considered suitable for affor- cone-bearing branchlets terete or 4-angled, 0.5–1.5
estation of deforested lands in its native area because mm long, with 6–8(–12) imbricate, obtuse scales.
of its easy germination and light-demanding proper- Seed cones subterminal, solitary or paired at apex
ties combined with rapid growth. Its wood has good of lateral branchlets, ovoid, (4–)5–6(–7) mm long,
properties, e.g. durability, but trees tend to be much (2.5–)3–4 mm wide, dehiscent when mature (in first
branched especially when grown in open vegetation. year), with 4 decussate, flattened scales (very rarely
It is less hardy than C. decurrens or C. formosana but with 2 additional, basal scale rudiments). Bract-
is in cultivation in parks and large gardens in China scale complexes somewhat coriaceous, green, be-
and elsewhere in SE Asia. coming woody and brown, broadly ovate, 4–6 mm
long, 2.5–4 mm wide; basal 2 scales fertile, dehiscent,
normally 2-seeded, rarely with 1 seed; apex rounded,
Calocedrus rupestris Aver., Hiep & L. K. Phan, Proc. incurved, sometimes with an indistinct, slightly flat-
Nat. Conf. Life Sci. Thai Nguyen Univ. Sept. 2004: tened or concave, subapical plate with a rugose sur-
41. 2004. Type: Vietnam: Bac Kan Prov., Na Ri face, rarely with a small central umbo (bract tip);
District, Na Bo, L. V. Averyanov et al. 5441 (holo­ apical pair of scales sterile, connate. Seeds ovoid or
type HN). Fig. 75 sub-ovoid, acute at apex, slightly flattened, including
the two relatively large but very unequal wings 4–5
mm long.
Etymology
The species epithet means ‘rock-dwelling’ and refers
Taxonomic notes
to the habitat in which it is invariably found.
Ongoing surveys of the rich flora of Vietnam result
in the frequent discovery of new taxa, new to the
Vernacular names
country or even to science, and this includes coni-
No common names have been recorded for this spe- fers. This is a new species in the genus Calocedrus,
cies. which seems to be restricted to the karst limestone
formations of northern Vietnam, already renowned
for its diversity and endemism. This species differs
Description
from C. macrolepis, also in Vietnam, mainly in the
Trees to 25 m tall; trunk to 1–1.2 m d.b.h. Bark 8–12 much smaller leaves and (consequently) cones, with
mm thick, fissured on large trunks, fibrous and ex- the seed cones having only two pairs of scales in-
foliating in longitudinal strips, resinous, greyish stead of three. The tree is not dwarfed and can attain
brown to brown. Branches spreading and ascending, considerable size, though due to growth conditions
forming a broadly rounded crown. Foliage branch- it is often small and even stunted. The two species
lets spreading and ascending, arranged in a plane, clearly occupy different habitats and are also geo-
covered with scale leaves, flattened, uniformly green graphically nearly separated.
or with very indistinct, whitened stomatal bands
on the underside. Leaves scale-like, decussate, in
 Distribution Conservation
Vietnam, N of the 17th parallel to the Chinese bor- This species is apparently more widespread than the
der. So far it has been found in seven provinces: recently discovered species Xanthocyparis vietnam-
Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Nghe An, ensis, also belonging to the Cupressaceae. It occupies
Quang Binh, and Son La. It may be present across the same or similar habitats and while locally com-
the borders of Laos and China (Yunnan). mon, in other regions it is rare. As with other mem-
TDWG codes: 41 VIE bers of the Cupressaceae in this part of the world, the
fragrant and durable wood is highly prized and log-
ging of the larger trees, even from inaccessible sites,
Ecology
is a problem for the conservation of the species. The 251
Calocedrus rupestris occurs on rocky limestone timber is marketed locally as well as across the bor-
mountains in northern Vietnam. These mountains der into China. Recruitment from seedlings is re-
are the eroded remains of a vast plateau of karst ported to be low, with very few seedlings observed
limestone that extends into China. This plateau is in most of the explored localities (Averyanov et al.,
in many places sharply dissected, leaving tower-like 2005). Due to deforestation and logging observed in
formations with steep, narrow ridges at altitudes many areas within the range and in the habitat of
from 600 m to 1600 m a.s.l. These ridges and summit this species, a serious reduction estimated at 50% or
areas bear a dense vegetation of small to medium- more in population size has been inferred in many
size trees, shrubs, epiphytes, ferns and mosses. Com- localities where this species has so far been found
mon conifers are Pinus kwangtungensis, Pseudotsuga by botanists. The total number of mature trees is es-
sinensis, Tsuga chinensis, Keteleeria davidiana (lo- timated to be below 2500. This species is still com-
cally), K. evelyniana (Pinaceae), ­Fokienia hodginsii, mon, with well structured age distributions in the
locally Xanthocyparis vietnamensis (Cupressaceae) population, within the limestone areas of Phong
and Amentotaxus hatuyensis, and more widespread Nha – Ke Bang National Park and in Bat Dai Son
A. yunnanensis (Taxaceae), Cephalotaxus ­mannii Nature Reserve.
(Cephalotaxaceae), Dacrycarpus imbricatus, Dacry- IUCN: EN (A2cd, C1)
dium elatum, Nageia fleuryi, and Podocarpus nerii­
folius (Podocarpaceae). There are many, mainly
Uses
small-leaved, broad-leaved trees (angiosperms)
e.g. Quercus spp., Acer tonkinense, Cinnamomum The wood of this species is valued for its fragrance
sp., Machilus sp., Carpinus sp., and Celtis sp., and and durability and is used for carpentry, interior fin-
epiphytic as well as lithophytic orchids. These karst ishing and furniture. This species, recently discov-
mountains are rich in endemic species and receive ered and described, is not yet in cultivation.
high levels of rainfall (1200–3000 mm p.a.) and fog.
 Cathaya Chun & Kuang, Acta Bot. Sin. 10 (3): 245. 1962. Type: Cathaya argyrophylla
Chun & Kuang (Pinaceae).

Cathay is an old name for (northern) China. bud scales ovate; basal scales smaller than the api-
cal scales, light yellowish brown, deciduous. Leaves
spirally arranged, on short shoots seemingly in tufts,
Description
spreading radially, densely crowded near shoot
See the species description. apex, directed forward, (1.8–)2.8–4.5(–5.5) cm long,
(2.2–)2.5–3 mm wide, more or less petiolate at base,
252 linear, flattened, longitudinally grooved and green
Distribution
above, with 2 glaucous white bands below; mar-
As for the species. gins slightly recurved, ciliate in young leaves; apex
obtuse. Stomata in two bands separated by a mid-
rib below, in ca. 15 rows on each side. Pollen cones
Cathaya argyrophylla Chun & Kuang, Acta Bot. lateral, near shoot apex, pendant, with involucre of
Sin. 10 (3): 246. 1962. Pseudotsuga argyrophylla light brown, large scales, 3–6 cm long, yellowish.
(Chun & Kuang) Greguss, Bot. Közlem. 57: 54. Seed cones lateral, at about right angles from shoot,
1970; Tsuga argyrophylla (Chun & Kuang) de Laub. short pedunculate, ovoid oblong, with acute apex,
& Silba, Phytologia Mem. 7: 75. 1984. Type: China: 3–5 cm long, 1.5–2.5 cm wide with slightly opened
Guangxi, Guangfu, [“ad regionem sylvaticam seed scales, greenish, maturing to orange brown,
Kwangfuensem”], Kwangfu Lingchü Exped 198 ripening to chestnut brown. Bracts triangular, with
(holotype IBSC, isotype PE). Fig. 76, 77 a long point, 5–8 mm long, included. Seed scales
13–16, orbicular or ovate, 1.5–2.5 cm long and 1–2 cm
wide at mid-cone; surface longitudinally striated or
Etymology
furrowed, initially puberulent, later glabrous; upper
The species epithet is derived from the Greek argyr- margin entire, soon becoming erose; base very short
= silver and phyla = leaf. pedicellate. Seeds obovoid, with an acutish base, 5–6
× 3–4 mm, dark olive with pale green spots; seed
wings adnate, obovate cuneate, 10–15 mm long and
Vernacular names
4–6 mm wide, light brown.
yin shan (Chinese)
Taxonomic notes
Description
Cathaya argyrophylla is one of the prime examples of
Trees to 20 m tall, d.b.h. to 40–60 cm; trunk mo- a relict conifer which had a very much wider distri-
nopodial, straight, columnar, sometimes forked in bution in the geologic past. In the Tertiary Cathaya
the top; crown pyramidal, often irregular, open. occurred in Europe, Russia, and Canada, and the
Bark grey, becoming scaly, lower down the trunk of genus is rather common as a fossil in the Miocene
older trees breaking into irregular plates. Branches lignite (‘brown coal’) deposits of Germany. Its taxo-
of first order long, heavy, spreading horizontally or nomic position has been the subject of some dispute
slightly ascending; branches of second order slender, in the past, but more recent phylogenetic studies in
branching irregularly. Branchlets slender, firm, yel- the Pinaceae have confirmed its status as a genus,
lowish, becoming grey in second year; surface with probably related to Pseudotsuga in a ‘pinoid’ clade.
ridges ending in weak pulvini, divided by grooves,
glabrous, but very young shoots puberulent; leaf
Distribution
scars angular circular; no real short shoots (weak
dimorphism), but alternately long and short growth China: Chongqing (Nanchuan Xian, Wulong Xian),
of shoots, the lateral (weaker) shoots forming 1–3 NE Guangxi (Jingxiu Yaozu Zizhixian, Longsheng
sequences with fewer and shorter leaves. Vegeta- Gezu Zizhixian), Guizhou (Daozhen Xian, Jiangkou
tive buds ovoid conical, 6–8 mm long, not resinous;
Xian, Tongzi), S Hunan (Chengdu, Guidong Xian, its extent of occurrence (EOO) is now known to en-
Luohandong, Xinning Xian, Zixing Shi). compass four provinces in south-central China and
TDWG codes: 36 CHC-CQ CHC-GZ CHS-GX CHS- herbarium collections (in China) are known from at
HN least 10 localities. It is usually growing on inacces-
sible slopes and ridges and is not considered to be
a valuable timber resource due to small or medium
Ecology
size and ‘poor shape’ in logging terms. Several lo-
On medium high mountains, at elevations between calities are within reserves and enjoy legal protec-
(900–)1200 m and 1900 m a.s.l. The soils are the tion. The species is now in cultivation in China and
widely distributed red and yellow earths of humid, is slowly becoming available through some botan-
warm temperate to subtropical China. Wang (1961) ic gardens in the West, e.g. Royal Botanic Garden 253
has mentioned C. argyrophylla as a rare conifer Edin­burgh. It is probably in the category Least Con-
occurring in the evergreen sclerophyllous broad- cern (LC) but a reassessment of the old status (under
leaved forest type. This forest type is dominated by IUCN criteria version 2.3, 1994) is overdue.
numerous species of Fagaceae with mostly small, IUCN: NT
ovate lanceolate, coriaceous leaves. However, from
the altitudinal range of the species it is likely that
Uses
it occurs in an ecotonal type between the sclero-
phyllous and deciduous broad-leaved forest types. This species is in cultivation through several forest-
Other conifers with which it occurs are almost cer- ry institutes and botanic gardens in China; outside
tainly ­Pinus fenzeliana (syn. P. kwangtungensis), and China it is still very rare in collections and no ma-
possibly also Tsuga chinensis and Nothotsuga longi­ ture plants exist in these. It has only recently been
bracteata. freed of its official embargo, which was not based in
considerations of conservation of a rare species but
in geopolitics. Lifting of these strictures may have
Conservation
begun with the donation of some seedlings by Prof.
After its discovery in the 1950s, for many years this Fu Likuo when he visited the Royal Botanic Garden,
monotypic genus was considered to be an extremely Edinburgh in 1998 for work on Volume 4 of the Flo-
rare conifer. Even herbarium specimens were very ra of China. Although the species is now available in
few and virtually nothing of it had reached botanic the trade in both Europe and the USA, it will prob-
gardens and institutional herbaria outside China ably remain a tree for collector’s gardens only, as it
until very recently. It is a relatively rare conifer, but has no remarkable horticultural merit.
 Cedrus Trew, Traité Arbr. Arbust. 1: 139. 1755 (nom. cons.). Type: Cedrus libani
A. Rich. (Pinaceae).

Cedrus is the classical Latin name for the (true) ce- s­ eparate (disjunct) regions. If the genus once had
dars; however, it was also in use for (fragrant) wood a more continuous distribution, with populations
of other conifers. bridging the great gaps now extinct, it is evident that
this fragmentation happened long ago. Such separa-
tion leads inevitably to genetic differences between
Description
the remnant populations, and eventually to sepa-
254 Monoecious evergreen trees with a columnar trunk. rate species. A comprehensive genetic (DNA based)
Resin canals in bark, leaves, and seed cones, resin study of the genus is necessary, but has not yet been
cysts in wood. Young trees branching at regular in- undertaken and published.
tervals, later several branches of the first order be-
come codominant with the main stem, except in 3 species.
C. deodara, where the apical dominance is main-
tained (Massart’s model). Bark on large trunks
Distribution
breaking into small, irregular plates and becoming
longitudinally fissured. Shoot dimorphism pro- Disjunct: Atlas Mountains of Morocco and Algeria;
nounced, with long, slender leading shoots and Lebanon, Syria, SE Turkey, Cyprus; Himalaya.
lateral, short spur shoots; a terminal bud forming a
long shoot, and several lateral buds forming short
Key to the species of Cedrus
shoots. Leaves linear-acicular, rigid, usually slightly
narrowed near base and tapering towards an acute 1a. Leading shoot and tips of branches drooping.
apex, amphistomatic, diamond shaped in cross sec- Leaves on short shoots 2–4.5 cm long. Pollen
tion. Pollen cones solitary and more or less erect cones usually longer than 5 cm C. deodara
from apex of short shoots, large, catkin-like; pol- 1b. Leading shoot and tips of branches erect or
len bisaccate; phenology marked by late shedding spreading. Leaves on short shoots less than 3
of pollen (September to November), related to late cm long. Pollen cones shorter than 5 cm 2
development of the female strobilus. Seed cones 2a. Seed cones when fully mature up to 12 cm long
similarly situated, distinctly erect, requiring about (but sometimes not exceeding 10 cm). Pollen
17–18 months for full development, becoming ovoid cones up to 5 cm long. Indigenous to E Medi-
to barrel shaped when full grown. Bracts small and terranean C. libani
hidden, not growing with the seed scales. Seed scales 2b. Seed cones when fully mature up to 8 cm long.
helically arranged on a narrowly conical rachis and Pollen cones up to 4 cm long. Indigenous to
extremely imbricate, large, flabellate, coriaceous or W Mediterranean (Algeria and Morocco)
thin woody, dismembering from the persistent ra- C. atlantica
chis by abcission at maturity. Seeds fully covered by
a membrane on one side and partly covered with a
small portion at other side, membrane continuing in Cedrus atlantica (Endl.) Manetti ex Carrière, Traité
a very large, broad seed wing. Seedlings with (5–)8– Gén. Conif.: 285. 1855. Cedrus libani A. Rich. var.
10(–14) cotyledons. atlantica (Endl.) Hook. f., [Cedars Lebanon] Nat.
Hist. Rev., ser. 2, 2: 15. 1862; Cedrus libani A. Rich.
subsp. atlantica (Endl.) Batt. & Trab., Fl. Algérie
Taxonomic notes
Tunisie: 397. 1905. Type not designated. Fig. 78
As will be discussed in more detail below with the
species, the morphology commonly deployed to
Etymology
circumscribe and differentiate the constituent spe-
cies of the genus Cedrus is on critical examination The species epithet means ‘from the Atlantic’ con-
not very convincing. The three (or four) species trasting the species with the cedars that grow in the
traditionally recognized occur, however, in vastly eastern Mediterranean.
 Vernacular names Taxonomic notes
Atlas cedar; Cèdre de l’Atlas (French) The taxonomic distinction between Cedrus atlantica
and C. libani is at best inconsistent and, when evalu-
ated critically based on the populations growing
Description
in the wild, probably non-existent. Much has been
Trees to 30–35(–40) m tall, but at higher elevations made of glaucousness of foliage, but this is based
usually lower, d.b.h. to 1.5–2 m; trunk usually mo- on trees selected for horticulture; glaucous as well
nopodial, columnar, massive, often forked above the as green leaved trees occur in Morocco as well as in
middle. Bark on trunk cracked and fissured, rough, Turkey, often in the same populations. Numbers of
dark grey, breaking into flaking plates revealing red- leaves on short shoots and length of leaves not only 255
dish brown bark. Branches of first order massive, vary, the values often cited as distinct between the
the upper ones ascending, the lower horizontal or two species can be found in both distant areas. The
bent downward; branches of second order crowded, smaller maximum size of seed cones in C. atlantica
spreading in horizontal or descending planes, as- may be correct, but to emphasize this hides the sub-
cending near the top of the tree; crown in young stantial overlap in cone sizes actually observed in
trees broadly conical, in old trees spreading later- natural populations. Crown shape is entirely a ques-
ally, becoming flat topped. Branchlets short, firm, tion of age and of growing conditions; roughly speak-
except leading shoot, which is longer and slender, ing: the older the tree the flatter the crown in both
grey green or grey brown, soon turning grey and species. In Turkey, trees with pyramidal crowns are
flaking, densely blackish pubescent at first; pul- known as C. libani subsp. or var. stenocoma, but that
vini on long shoots small, prominent; short shoots distinction is taxonomically doubtful, too. Should
thick, scaly, of variable length with age (0.5–3 cm), we therefore unite the Mediterranean Cedars under
assurgent or erect. Vegetative buds ovoid globular, a single species, C. libani? Should we recognize sub-
2–3 × 1.5–2 mm, not resinous; bud scales broadly species or varieties? The only true distinction is the
ovate, red brown, dark brown or blackish at apex, great geographical separation, which rules out gene
deciduous. Leaves on long shoots spirally arranged, flow between C. atlantica and C. libani in nature.
remote, near base of long shoot more crowded, ra- They may become truly distinct species in the future.
dially spreading, falling in 2nd or 3rd year; on short
shoots densely crowded in false whorls, 20–45,
Distribution
spreading radially, (1–)1.5–2.5(–3) cm long, 1–1.5
mm wide, narrowly linear, straight or curved, dia- NW Africa: Algeria, Morocco (Atlas Mts.).
mond shaped in cross section; apex acute or acumi- TDWG codes: 20 ALG MOR-MO
nate; stomata on all sides, more numerous on two
adjacent sides; leaf colour glossy dark green or glau-
Ecology
cous. Pollen cones terminal on short shoots, erect,
subtended by leaves, numerous, soon falling after The Atlas cedar occurs on the high maritime ranges
shedding pollen, 3–4 cm long, straight, later curved, of the Atlas Mountains, at elevations between 1370 m
first rose yellow, later pale brown. Seed cones termi- and 2200 m a.s.l., especially on N and NW exposed
nal on short shoots, erect, sessile, becoming woody slopes receiving 1000 to 2000 mm precipitation an-
in 2nd year, ovoid or barrel shaped; apex obtuse or nually, mostly during the winter. At the high ridges
retuse, 5–8 × 3–5 cm, light green, maturing to pale much snow accumulates. The summers are warm
green, with purplish edges of seed scales, ripening and dry. The soil is usually rocky, calcareous and well
to light (purplish) brown; cone rachis persistent, drained. The species grows in pure stands, reach-
narrowly conical. Seed scales broad flabellate, thin, ing tree limit on high, exposed ridges, or is locally
coriaceous, length × width 2–3 × 2.5–3.5 cm; surface mixed with Abies numidica, Taxus baccata, Quercus
smooth, orange-brown pubescent at base, glabrous faginea, Acer obtusatum, Populus tremula, Sorbus
on exposed parts; upper margin entire, slightly in- aria, and S. torminalis (Algeria) or Abies pinsapo
curved; base pedicellate. Seeds ovoid conical, 8–13 × var. marocana, Juniperus communis, Taxus baccata,
4–6 mm, brown; seed wings broad cuneate, 18–25 × Populus tremula, Quercus ilex, and Acer granatense
12–17 cm, (light) brown. locally in Morocco.
 ing in horizontal or descending planes, the lead-
Conservation
ing shoots drooping; crown of young trees conical,
IUCN: LC with drooping leader, old trees conical. Branchlets
short, firm, but leading shoots long and slender, pale
(pinkish) fawn brown, becoming grey brown; sur-
Uses
face smooth, soon breaking, showing green bark in
Atlas cedar is an important tree for timber in North cracks, densely light brown pubescent at first; pul-
Africa, providing strong and durable wood for con- vini small, prominent on long shoots; short shoots
struction purposes as well as for the manufacture thick, scaly, of variable length with age (0.5–4 cm),
of furniture. In the 19th century it was introduced assurgent or erect. Vegetative buds ovoid, acute, 1–2
256 in Europe (France, 1839) and rapidly spread as an mm long, not or slightly resinous; bud scales ovate,
ornamental tree desirable for landscape gardens, of- orange brown, with pale brown apex, deciduous.
ten supplanting the much earlier introduced Leba- Leaves on long shoots spirally arranged, remote,
non cedar. Despite claims to the contrary, which are more crowded at base of long shoot, falling after
mostly based on perceptions, not on knowledge of 2–2.5 years; buds of short shoots axillary; leaves on
characters of natural populations, the two putative short shoots densely crowded in false whorls, 20–30,
species are extremely difficult, if not impossible, to spreading radially, 2.5–4.5 cm long (2–6 cm on long
distinguish as mature trees in cultivation. A very shoots), 1–1.5 mm wide, narrowly linear, straight or
glaucous leaved form is particularly in demand for slightly curved, diamond shaped in cross section,
horticulture and is based on selections (‘Glauca’ or abruptly acute; more or less amphistomatic, but
‘var. glauca’) of individuals found in nature. Several stomata on two adjacent sides more numerous; leaf
other cultivars are known, based on habit and foli- colour green to grey green. Pollen cones terminal on
age characteristics. short shoots, erect, subtended by leaves, numerous,
soon falling after shedding pollen, (4–)6–7 cm long,
immature greenish, ripening to rose brown. Seed
Cedrus deodara (Lamb.) G. Don, in Loudon, Hort. cones terminal on short shoots, erect, sessile, be-
Brit. 1: 388. 1830. Pinus deodara Lamb., Descr. Pinus coming woody in 2nd year, usually barrel shaped or
2: 8. 1824; Cedrus libani A. Rich. subsp. deodara ovoid; apex obtuse or retuse; 7–13 × 5–9 cm, pale grey
(Lamb.) P. D. Sell, Watsonia 18 (1): 92. 1990. Type green, maturing to glaucous green, sometimes with
not designated. Pl. 10, Fig. 79 purple, ripening to dark brown; cone rachis persist-
ent, narrowly conical. Seed scales broad flabellate
or rectangular flabellate, thin, coriaceous, length
Etymology
× width 3–4 × 3.5–4.5 cm; surface smooth, yellow-
Deodar is the vernacular name for this species in ish brown pubescent at base; upper margin entire,
India. thin, slightly incurved; base pedicellate. Seeds ovoid
conical, 10–15 × 5–7 mm, brown; seed wings broad
cuneate, 20–30 × 15–21 mm, light brown.
Vernacular names
Deodar cedar, Himalayan cedar; deodar (India);
Taxonomic notes
xuesong (Chinese)
Reducing Cedrus deodara to a subspecies of C. li-
bani, as was proposed by Peter Sell (op. cit.), would
Description
presumably reduce Cedrus Trew to a monotypic
Trees to 40–50(–65) m tall, d.b.h. to 2.5–3 m; trunk species, with only a few varieties or subspecies rec-
usually monopodial, straight, columnar, sometimes ognized under that species. Based on morphology,
forked in several stems above 2/3 of bole. Bark soon there is merit in this view, but it may not be the en-
cracked and flaking, breaking into small plates, fis- tire truth about what in this case actually constitutes
sured, dark blackish grey. Branches of first order a distinct species. Unlike between C. atlantica and
massive, spreading horizontally or almost erect C. libani, there are consistent, albeit small, morpho-
near the top, the lower branches curved downward; logical differences between these two species and
branches of second order crowded, usually spread- C. deodara. More research into genetic aspects of the
 6
257

2 5

Pl ate 10. Cedrus deodara. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone. 4. Pollen cone and
leaves. 5. Leaf. 6. Seed.
 problem is needed before such a far reaching con- are commonly used in gardens. This use has led to
clusion can be drawn. the selection of several cultivars, with different habit
including dwarf forms and/or with varying foliage
colours; most of these are in cultivation in Central
Distribution
Europe.
E Afghanistan (Hindu Kush), NW Pakistan (Kara-
koram), China: extreme SW Xizang [Tibet], Kash-
mir to W Nepal. Cedrus libani A. Rich., in Bory, Dict. Class. Hist.
TDWG codes: 34 AFG 36 CHT 40 NEP PAK WHM- Nat. 3: 299. 1823.
HP WHM-JK WHM-UT
258
Etymology
Ecology
The species epithet denotes the Lebanon from where
Cedrus deodara is a high mountain tree, but it oc- this species was first described and named.
curs in a wide range of habitats in the Himalaya. It
grows in a belt at elevations between 1700 m and
Vernacular names
3000 m a.s.l. in the western part of its range and be-
tween 1300 m and 3300 m in the eastern part, where Cedar of Lebanon, Lebanon Cedar
the climate is less dry. It grows on a variety of alpine
lithosols. The climate is moist monsoon, but the in-
Description
creasing moisture in the E Himalayas is a limiting
factor; towards the west it becomes moderately dry, Trees to (20–)35–40 m tall, d.b.h. to 1.5–2.5 m; trunk
with annual precipitation less than 750 mm in the usually monopodial, columnar, massive, often
most western part of its range. At higher elevations forked or branched below 1/3 of the bole. Bark soon
it forms a coniferous forest belt with, among other cracked and scaly, rough, with flaking small plates
species, Abies pindrow, A. spectabilis in Nepal, Pinus and deep, longitudinal fissures, dark grey to blackish
wallichiana, Picea smithiana, and Cupressus torulo- brown. Branches of first order massive, often erect,
sa, but forms often also pure stands. At the highest eventually spreading horizontally, the lowest often
limits of Cedrus, Juniperus squamata is the only ac- weighed down at ends; branches of second order
companying conifer species. At lower elevations first crowded, spreading in horizontal planes; crown in
Quercus spp., then Aesculus indica, Betula sp., Cory- young trees conical (sometimes retaining a narrow
lus jaquemontii, Acer spp., Prunus spp. and shrubs habit longer), but later spreading horizontally, often
mark the transition towards a broad-leaved forest. several boles forming crowns at different levels, flat
topped, but in forest stands more pyramidal. Branch-
lets short, firm, but leading shoot long and slender,
Conservation
(pale) brown, turning grey, shallowly grooved, soon
IUCN: LC flaking, at first with dark brown pubescence; pul-
vini small, on long shoots; short shoots thick, scaly,
of variable length with age (0.5–4 cm), assurgent
Uses
or erect. Vegetative buds ovoid, 2–3 × 1.5–2, not or
Himalayan cedar is a very important timber tree in slightly resinous; bud scales broadly ovate, free at
Pakistan, Kashmir and NW India. Its strong and du- tips, pale brown with dark apex, deciduous. Leaves
rable wood is mostly used for construction. Other on long shoots spirally arranged, remote, radially
uses are general carpentry and furniture. A fragrant spreading, more crowded at base of the long shoot;
oil can be distilled from wood chips and sawdust. short shoots emerging in leaf axils, leaves densely
It was first introduced in England in 1822 as an crowded in false whorls, (15–)20–35, most numerous
ornamental tree, but as a park tree it remains less on older short shoots, spreading radially, (0.5–)1–2.5
common than the two Mediterranean species. It is cm long (on long shoots 1–3.5 cm), 1–1.5 mm wide,
generally more susceptibe to ‘late’ frost than these narrowly linear, straight or curved, in cross sec-
and also requires more moisture. Young trees espe- tion diamond shaped, acute or acuminate; more or
cially have a distinct habit with drooping ­leaders and less amphistomatic; leaf colour light or dark green,
sometimes glaucous. Pollen cones terminal on short and locally J. foetidissima, at lower elevations it is re-
shoots, erect, subtended by leaves, numerous, 4–5 placed by Pinus nigra and Pinus brutia.
cm long, cylindrical, curved when ripe, pale green
at first, ripening to pale brown. Seed cones terminal
Uses
on short shoots, erect, sessile, becoming woody in
2nd year, ovoid, ovoid oblong or barrel shaped, with Cedar of Lebanon is a timber tree used since the
obtuse or retuse apex, (5–)8–12 × 3–6 cm, first light dawn of civilization. About 4600 years ago King
green, maturing to greyish or purplish green, ripen- Snefru of the Fourth Dynasty of ancient Egypt, who
ing to grey brown with purplish edges of seed scales; ordered the building of the pyramids of Dahsjur
cone rachis persistent, narrowly conical. Seed scales and Meidum, used 40 shiploads of cedar timber for
broad rectangular flabellate, thin, coriaceous, soon shipbuilding and for the heavy doors of the king’s 259
breaking up below, length × width 3–3.5 × 3.5–4 cm; palace. It was undoubtedly used in similar ways be-
surface smooth, orange-brown pubescent at base; fore this earliest written record. Solomon’s temple in
upper margin entire, thin, slightly incurved; base Jerusalem is reputed to have been built with its tim-
pedicellate. Seeds ovoid conical, 10–14 × 4–6 mm, bers. Subsequent civilizations plundered the Leba-
brown; seed wings broad cuneate, 20–30 × 15–18 non cedar forests to almost depletion. A tree of this
mm, light brown. species appears as a symbol in the flag of Lebanon.
In Turkey, where it is much more abundant, forests
are now mostly well managed to provide a renew-
Taxonomic notes
able resource of strong, hard, and durable timber
The variety C. libani var. stenocoma, in earlier bo- for construction, boat building and furniture. It was
tanical literature usually recognized as a sub­species introduced in Europe as early as 1638 (England) and
and said to be intermediate between Cedrus atlan- became a major tree in the landscaped parks around
tica and Cedrus libani, is probably more accurately stately homes in much of Europe, but especially in
merely an ecotype. Given variability of morphol- Britain. Many trees at these locations are now majes-
ogy in C. libani (and C. atlantica) among trees tic specimens, often with multiple stems and huge,
with spreading to flat-topped crowns, there are no spreading flat crowns. Cultivars have been selected,
characters that differ in this variety, and we are left but are less numerous than those derived from Atlas
with its habit. As it often occurs in relatively dense and Himalayan cedars.
stands mixed with Abies cilicica or in pure stands of
relatively young trees, it is probably assuming this 2 varieties are recognized:
growth form due to competition. Such trees would
be understood as ecotypes and not as proper taxa. Cedrus libani A. Rich. var. libani. Type: Illustration
of “Cedrus” in Belon, Plur. Rer. Obs.: 162. 1605
(lectotype). Fig. 80
Distribution
Lebanon, Syria, Turkey. Cedrus libanitica Trew ex Pilg. subsp. stenocoma
TDWG codes: 34 LBS-LB LBS-SY TUR O. Schwarz, Fedde’s Repert. Sp. Nov. Regni Veg. 54
(1): 26. 1944; Cedrus libani A. Rich. subsp. stenocoma
(O. Schwarz) P. H. Davis, J. Roy. Hort. Soc. London
Ecology
74: 113. 1949; Cedrus libani A. Rich. var. stenocoma
In high mountains around the eastern Mediterra- (O. Schwarz) Frankis, Fl. Turkey & E. Aegean Is. 11:
nean basin, especially on N-facing slopes, at eleva- 5. 2001.
tions between 1300 m and 3000 m a.s.l. The soils are
usually well drained, calcareous lithosols on rocky
Description
slopes and ridges. The climate has cool, moist win-
ters (annual precipitation 1000 mm to 1500 mm) Trees pyramidal when young, older trees develop-
and warm, dry summers; at the highest elevations ing broad and eventually flat-topped crowns. Leaves
there is abundant snow in winter. Cedrus libani oc- (1–)2–2.5(–3.5) cm long, green, grey-green, or glau-
curs in pure stands or mixed with Abies cilicica in cous green.
Turkey, in which are also common Juniperus excelsa
 not show the same variation as is found in natural
Distribution
populations; taxa should always be circumscribed
Lebanon, Syria (Djebel el Ansiriya), Turkey (Taurus according to characters observed in wild popula-
and Anti-Taurus Mts., disjunct in northern Turkey). tions, if possible. It can be maintained as a variety
TDWG codes: 34 LBS-LB LBS-SY TUR but merely on the basis of a trait: often shorter leaves
than in C. libani var. libani.
Conservation
Distribution
IUCN: LC
Cyprus, Troodos Mountains, Mt. Triphylos.
260 Cedrus libani A. Rich. var. brevifolia Hook. f., TDWG codes: 34 CYP
J. Bot. 18: 31. 1880. [J. Linn. Soc., Bot. 17: 518. 1880]
Cedrus brevifolia (Hook. f.) A. Henry, in Elwes &
Ecology
Henry, Trees Gr. Brit. Ireland 3: 467. 1908; Cedrus
libani A. Rich. subsp. brevifolia (Hook. f.) Meikle, Cedrus libani var. brevifolia occurs on medium high
Fl. Cyprus 1: 22. 1977. Type: Cyprus: Southern mountains at elevations between 900 m and 1525
Cyprus, Troödos Mts., Troödos Forest, S. Baker s.n. m a.s.l. in two separated populations on N-facing
(holotype K). slopes, where the soils are calcareous and rocky. The
climate is mild and wet in winter, with ca. 1000 mm
annual precipitation, and warm and dry in summer.
Description
The two almost pure, scattered stands have been
Trees to 15–20 m tall, d.b.h. 1–1.2 m, eventually flat- much degraded by cutting, fire and goats and are
topped. Leaves 0.5–1.6(–2) cm long, (sometimes only remnants of formerly more extensive forests.
longer in cultivation), the longest at base of long
shoots, width 1–1.5 mm.
Conservation
This variety of C. libani has a limited distribution
Taxonomic notes
and is known from two populations in the moun-
This taxon, although often recognized as a distinct tains of Cyprus. The geographical isolation on the is-
species, is on critical examination of doubtful dis- land of Cyprus could have led to genetic distinctions
tinction. Many trees appear to have very short leaves, not expressed in the morphology; if so we would be
but some (and not only young trees) have leaves as dealing with a threatened genotype isolated from its
long as those found on many trees of var. libani in main congeners. However, the same would probably
Turkey, Syria and Lebanon. Also, in var. libani forms be true of the remnant population of var. libani in
with short leaves can be found in Turkey, and if the mountains of Lebanon, further (and perhaps
short leaves are the only distinctive character, var. longer) disjunct from the main range of the species
brevifolia could be more widespread or both varie- than that of Cyprus, but in which no morphological
ties not taxonomically distinct. Trees taken into cul- distinctions are discernible.
tivation may be of limited provenance and therefore IUCN: VU (D2)
Cephalotaxus Siebold & Zucc. ex Endl., Gen. Pl. Suppl. 2: 27. 1842. Type: Cephalo­
taxus harringtonii (Knight ex J. Forbes) K. Koch (Taxus harringtonii Knight ex
J. Forbes) [Cephalotaxus pedunculata Siebold & Zucc. ex Endl. (nom. illeg.)]
(Cephalotaxaceae).

Key to the species of Cephalotaxus

Greek: kephalos = head; referring to the structure
(cone) onto which the seed is attached; Taxus is the The number of lines of stomata in a stomatal band
classical Latin name for yews. is not necessarily correlated with the width of that
band, so a magnifying lens 10–20 × is required to 261
observe this character. The disposition of leaves (in
Description
a horizontal plane or in a V-formation) can vary
Dioecious evergreen shrubs or small trees. Resin between fertile and infertile shoots and between
canals (1) in the leaves only. Bark thin, smooth, ex- sun-exposed and shaded shoots and is by itself not a
foliating in narrow strips. Branches spreading and reliable character.
plagiotropic or ascending, foliage branches termi-
nating in conical, scaly buds. Leaves spirally in- 1a. Leaves convex, touching each other closely,
serted, spreading in all directions on erect shoots, cordate or truncate at base; apex minutely cus-
pectinately arranged on lateral, plagiotropic shoots pidate or mucronate C. oliveri
by alternately twisting of petiolate leaf bases, becom- 1b. Leaves more or less flat, separately disposed,
ing subopposite, flattened, linear-lanceolate, with cuneate to nearly truncate or twisted petiolate
two prominent stomatal bands separated by a raised at base; apex acuminate or cuspidate, some-
midrib on the abaxial side. Pollen cones aggregated times mucronate 2
in pendulous, more or less globose, short-stalked 2a. Leaves mostly 3.5–10(–12.5) cm long, curved
clusters (capitulae) in the axils of leaves, forming op- upward and outward or downward, forming a
posite or subopposite rows on the underside of lat- V-formation. Peduncle of seed-bearing struc-
eral foliage branchlets; each small cone consiting of ture variable in length, but up to 20–25 mm
a short rachis with up to 15 peltate microsporophylls long 3
in spiral arrangement, each with 2–4 pollen sacs 2b. Leaves mostly 2–5(–7) cm long, spreading hori-
containing spherical pollen. Seed cones in clustered zontally or forming a V-formation on shoots
pairs or groups from axillary buds situated distally with pollen cones. Peduncle of seed bearing
from lateral shoots, (long) pedunculate, consisting structure up to 10(–15) mm long 4
of two sterile and several decussate fertile bracts, 3a. Leaves (1.5–)3–5 mm wide. Seed-bearing struc-
the fertile bracts with two axillary, erect ovules, tures 3–6 together; seeds including the striated
somewhat swollen at pollination and at that time or ridged aril 14–25 × 9–15 mm C. fortunei
still lacking an arillus, developing much later. Seeds 3b. Leaves 4–7 mm wide. Seed-bearing structures
from 1–2 fertilized ovules per cone, large, obovoid to solitary; seeds including the indistinctly stri-
ellipsoid, surrounded by a fleshy, green aril ripening ated aril 35–45 × 20–25 mm C. lanceolata
from green to yellowish, reddish or purplish brown. 4a. Stomatal bands with 10–15 lines of stomata; leaf
Seed proper large, with a hard, sclerified seed coat. bases cuneate 5
4b. Stomatal bands with 20–25 lines of stomata;
8 species. leaf bases obtuse to nearly truncate 6
5a. Pollen cones 6–7 per capitulum, to 3 mm diam.
Seeds including the aril ellipsoid, with 6 longi-
Distribution
tudinal ridges C. sinensis
NE India (Arunachal Pradesh, Assam); Myanmar 5b. Pollen cones 6–12 per capitulum, 3–4 mm
[Burma]; China; Korea; Japan; Taiwan; Laos; Thai- diam. Seeds including the aril obovoid, smooth
land; Vietnam. or striated 7
 6a. Leaves spreading at 45–80° to shoot axis, linear midrib and ­bordered by green, slightly revolute leaf
and abruptly narrowing to a mucronate apex margins. Pollen cones situated in rows of capitula
C. hainanensis on the underside of lateral foliage branchlets; each
6b. Leaves spreading at 70–90° to shoot axis, wid- capitulum on a 3–7 mm long, scaly peduncle insert-
est below the middle and gradually tapering to ed in the upper axil of a leaf, subtended by ovate,
a cuspidate apex C. mannii incurved bracts with entire or erose, more or less
7a. Leaves on shoots with pollen cones forming a hyaline margins, bearing 6–14 small, sessile or short
V-formation. Peduncle of seed-bearing struc- pedunculate, globose cones 3–4 mm diam. Micro-
ture 5–15 mm long C. harringtonii sporophylls 6–16 per cone, each with 3–4 globose,
7b. Leaves on shoots with pollen cones (as other cream coloured pollen sacs. Seed cones borne 3–6
262 leaves) spreading more or less horizontally Pe- together at base of lateral foliage branchlets, on
duncle of seed-bearing structure 2–5 mm long 5–25 mm long, slender peduncles. Bracts several
C. latifolia per cone, reduced, 1–1.5 mm long; fertilized ovules
surrounded by an ellipsoid, green or yellow aril, en-
closing the ripening seed, becoming 14–25 × 9–15
Cephalotaxus fortunei Hook., Curtis’s Bot. Mag. 76: mm, turning soft, purple and indistinctly striated or
t. 4499. 1850. longitudinally ridged, with a short mucronate apex.
Seeds ellipsoid or sometimes globose, 13–24 × 7–14
mm.
Etymology
This species was named after Robert Fortune, who
Distribution
introduced it to England in 1848.
SW, Central and SE China; N Myanmar [Burma].
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
Vernacular names
SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS-
Fortune’s plum yew; san jian shan, yuan bian zhong HE CHS-HK CHS-HN CHS-JS CHS-JX CHS-ZJ 41 MYA
(Chinese)
Ecology
Description
Cephalotaxus fortunei occurs as an understorey
Shrubs or trees 1–20 m tall; trunk to 40 cm d.b.h. tree or shrub in mixed broad-leaved (angiosperm)
or more, often multi-stemmed. Bark thin, exfolia- forests, in mixed conifer-broad-leaved forests and
ting in strips, reddish brown. Branches spreading or in coniferous forests (Abies and Picea). It is also
ascending, forming a broad crown in trees; shrubs common as a shrub or small tree in open thick-
often as wide or wider than tall. Foliage branchlets ets and on roadsides in secondary vegetation. Its
slender, terete, glabrous, finely grooved between de- altitudinal range is great and extends from 200 m
current leaf bases, green turning yellowish to light to 3700 m a.s.l., with C. fortunei var. alpina at the
brown. Leaves mostly spreading in two rows in a higher range between (1100–)1800 m and 3700 m
semi-pectinate arrangement, diverging at various a.s.l. In the Lower Yangtze Valley var. fortunei oc-
angles between 30–110° to shoot axis, commonly curs in remnants of mixed mesophytic forest with
curved upward and down towards apex; (1.5–)3.5– Acer spp., Catalpa ovata, Fraxinus chinensis, Ilex lati-
10(–12.5) cm long, linear-lanceolate, straight or folia, Liquid­ambar formosana, Nyssa sinensis, Quer-
slightly falcate, (1.5–)3–5 mm wide, with a short, cus spp., and many other angiosperm trees. Most of
twisted petiolate base, gradually tapering to an acute these forest remnants are disturbed and/or replaced
to cuspidate apex, coriaceous but flexible, dark dull by secondary shrubby vegetation, in which C. fortu-
green above, with two whitish green bands below. nei may recur. In the evergreen broad-leaved forests
Midrib on the adaxial (upper) side in a shallow de- of Fujian, which are also remnants, C. fortunei var.
pression, 0.5 mm wide, obtusely raised, continuous fortunei occurs in the understorey of oaks (Quercus
from base to apex, obtusely raised and continuous spp.) but mixed with many other tree species, some
on the abaxial side. Stomata in two broad bands of of which are conifers like Nageia nagi, Keteleeria for-
17–24 intermittent white lines separated by a green tunei and Fokienia hodginsii. In S Gansu, Shaanxi
and Sichuan var. alpina occurs in very different for-
Conservation
ests dominated by Abies, Picea or Larix, or a mixture
of these conifers, between 1800 m and 3600 m a.s.l. IUCN: NT
Here it is a subcanopy tree or shrub, often accompa-
nied by Taxus chinensis and shrubs like Eurya and Cephalotaxus fortunei Hook. var. alpina H. L. Li,
Rhododendron. Lloydia 16 (3): 164. 1953. Cephalotaxus alpina
(H. L. Li) L. K. Fu, Acta Phytotax. Sin. 22 (4):
282. 1984. Type: China, Yunnan, J. F. Rock 11572
Uses
(Holotype loc. ?)
The wood of this species is of limited economic val-
ue; the main uses are in horticulture. It grows into a 263
Description
large shrub or small tree and the long leaves of this
species are decorative. It is suitable for pruning and Leaves 1.5–3.5(–4) mm wide. Pollen cone capitula
hedge clipping. The hardiness will much depend sessile or very short pedunculate, peduncle 0–2 mm
on provenance, given the wide range of the species long. Aril covering seed with longitudinal ridges.
in China. A limited number of cultivars, some of
doubtful validity and originally described as varie-
Distribution
ties with shorter or longer leaves “than the type”, are
known and may still be in cultivation. Long-leaved China: S Gansu, Sichuan, N Yunnan.
forms seem to be especially popular with gardeners TDWG codes: 36 CHC-SC CHC-YN CHN-GS
and the Hillier Nurseries in Hampshire, England,
have developed several cultivars with such leaves in
Ecology
recent years. In China oil extracted from the succu-
lent aril enclosing the seed was traditionally used as An understorey shrub or small tree in mixed coni-
lamp oil. fer forests, e.g. with Abies spp., and along mountain
streams. Elevation (from GIS) 885–2554 m, from
2 varieties are recognized: herbarium specimens 1100–3520 m a.s.l.

Cephalotaxus fortunei Hook. var. fortunei. Type:

Conservation
China: [“200 miles N of Shang-see”], R. Fortune s.n.
(holotype K). Fig. 81, 82 The range of this variety indicates LC and no threats
or decline are known to occur. Probably confined
Cephalotaxus fortunei Hook. var. globosa S. Y. Hu, to the higher parts of SW China, true range may
Taiwania 10: 28. 1964. be poorly known due to difficulty of identification
and relatively inconspicuous habit. Extent of occur-
rence (EOO) based on herbarium specimens calcu-
Description
lated as 225,117 km² and area of occupancy (AOO)
Leaves (2.2–)3.5–5 mm wide. Pollen cone capitula on as 93,342 km² but probably larger. This variety is not
3–7 mm long peduncles. Aril covering seed with nu- known by its herbarium collections from protected
merous indistinct striations. areas, although some collections were made near the
Three Parallel Rivers Protected Area in Yunnan and
the variety may occur there, too.
Distribution
IUCN: LC
China: S Anhui, Chongqing, Fujian, S. Gansu, N
Guangdong, N Guangxi, Guizhou, SW Henan, W
Uses
Hubei, Hunan, Jiangxi, S Shaanxi, Sichuan, Yunnan,
Zhejiang; N Myanmar [Burma]. An ornamental shrub in horticulture.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS-
GX CHS-HE CHS-HN CHS-JS CHS-JX CHS-ZJ 41 MYA
 Cephalotaxus hainanensis H. L. Li, Lloydia 16 (3): soft, red with longitudinal striation. Seeds (ob)ovoid
164. 1953. Type: China: Hainan Island, [“Fan Yah” = to ellipsoid, sometimes laterally compressed, 18–28
Fanyang?], N. K. Chun & C. L. Tso 44183 (holotype × 8–14 mm, with a mucronate or cuspidate apex.
US).
Taxonomic notes
Etymology
In Flora of China 4: 87 (1999), this species is treated
The species epithet derives from Hainan, the island as a synonym of Cephalotaxus mannii. However, in a
(and now province) of China where this species is note the co-author Robert Mill considered the plants
native and perhaps indigenous. from Hainan to be separable as the species named
264 and described by H. L. Li, i.e. C. hainanensis. Mill
gives a statement of morphological differences be-
Vernacular names
tween the two species. Under this narrower species
Hainan plum yew; hai nan cu fei (Chinese, includes circumscription it is possible that other specimens
C. mannii in Flora of China 4, 1999). from southern China and northern Vietnam would
have to be included under C. hainanensis, with con-
sequences for its distribution and hence conserva-
Description
tion status. Ying et al. (2004) map this species as
Trees to 20 m tall; trunk to 70 cm d.b.h. Bark thin, occurring in W Guangdong, Hainan, SE and W
exfoliating in small or large flakes and strips, light Guanxi, W Yunnan, and even SE Xizang [Tibet]. The
brown to reddish brown, weathering grey. Branches populations of Yunnan and Tibet are here treated as
spreading or ascending, forming a rounded or nar- belonging to C. mannii. A critical modern revision of
row crown. Foliage branchlets up to 15 cm long, the genus to resolve these issues is lacking at present.
slender, grooved between decurrent leaf bases, green
turning orange-brown. Leaves arranged in two pec-
Distribution
tinate rows, spreading horizontally and at 45–80° to
shoot axis, subopposite, (1.5–)2–4 cm long, slightly China: Hainan Island (Jianfeng Ling, Limu Ling,
falcate, 2.5–4 mm wide, flat, base very short petio- Wuzhi Shan).
late or sessile, broadly obtuse to nearly truncate, leaf TDWG codes: 36 CHH
blade linear and abruptly narrowing to a mucronate
apex. Midrib thin but prominent and continuous
Ecology
from base to apex on the adaxial (upper) side, 0.3
mm wide, conspicuous but more flattened on the Cephalotaxus hainanensis occurs in mixed warm
abaxial side; leaf colour dark green or olive-green temperate to subtropical rainforests in mountainous
above, two white or glaucous white bands below. areas of the island of Hainan, where it attains tree
Stomata in two broad bands of ca. 20 intermittent, habit and size (10–20 m). Nothing has been record-
white lines, separated by the green midrib and bor- ed in non-Chinese literature about associated spe-
dered by slightly revolute leaf margins. Pollen cones cies at present, presumably these forests are diverse.
situated in rows of capitula on the underside of lat-
eral foliage branchlets; each capitulum on a (1–)3–5
Conservation
mm long, scaly peduncle inserted in the upper axil
of a leaf, subtended by ovate, incurved bracts with Under the narrower circumscription accepted here
entire or erose, more or less hyaline margins, bear- this species is probably restricted to Hainan but its
ing 6–8 small, sessile, globose, cream or light yellow entire range remains uncertain. It is threatened by
cones up to 4 mm diam. Microsporophylls 6–12 per timber harvesting and especially stripping of bark.
cone, each with 3–4 globose, cream coloured pol- IUCN: EN (A2d)
len sacs. Seed cones solitary at base of lateral foliage
branchlets, on 4–10 mm long peduncles. Bracts few
Uses
per cone, reduced, 1–1.5 mm long; fertilized ovules
surrounded by an ellipsoid, green aril, enclosing the The bark of this and other species is stripped to
ripening seed, becoming 20–30 × 10–15 mm, turning be used for medicinal purposes in China. It is not
 known to be in cultivation outside a few botanic ­ icrosporophylls 5–15 per cone, each with 2–3 glo-
M
­gardens. bose, cream coloured pollen sacs. Seed cones borne
3–6 together at base of lateral foliage branchlets, on
5–15 mm long, slender peduncles with small scale
Cephalotaxus harringtonii (Knight ex J. Forbes) leaves. Bracts several per cone, reduced, 1–1.5 mm
K. Koch, Dendrol. 2 (2): 102. 1873. long; fertilized ovules surrounded by an obovoid,
green or purplish aril, enclosing the ripening seed,
becoming 15–20(–25) × 12–18 mm, turning soft,
Etymology
orange-red to purple and smooth or striated, with
This species commemorates the Earl of Harrington, a short mucronate apex. Seeds (ob)ovoid to sub­
who gave a specimen to the Duke of Bedford, from globose, 12–18 × 8–12 mm. 265
whose pinetum at Woburn Abbey James Forbes de-
scribed and named it.
Taxonomic notes
This species has long been known as Cephalotaxus
Vernacular names
drupacea Siebold & Zucc. and it was introduced to
Harrington’s plum yew, Japanese plum yew; Inu- cultivation in the Netherlands by Von Siebold un-
gaya (Japanese); Picha-nam (Korean) der that name. However, this name was only validly
published in 1846, by which time the same species
(introduced by Von Siebold as stated) had been
Description
named and published as Taxus harringtonii [“har-
Shrubs to small trees 0.5–10 m tall; trunk d.b.h. of ringtonia”] by James Forbes in 1839, based in part on
monopodial trees to 40 cm. Habit prostrate to erect a manuscript received from the nurseryman Joseph
and spreading; trees with a wide, open and rounded Knight. It was later transferred to Cephalotaxus by
crown. Bark thin, exfoliating in narrow strips, grey- Karl Koch. Cephalotaxus koreana Nakai was de-
brown. Branches spreading, drooping or ascend- scribed as a 1–1.5 m tall, caespitose but non-layer-
ing, or prostrate in one variety; foliage branchlets ing shrub with “red [and] most delicious palatable
short, stout or slender, glabrous, grooved between fruits.” Those of C. drupacea (= C. harringtonii) were
decurrent leaf bases, green turning orange-brown. said to be bitter with a disagreeable smell. Otherwise
Leaves more or less disposed in two ranks, often up- the two taxa are similar and the assertion about ed-
right forming a V-formation especially on branch- ibility apparently being based on the experience and
lets with male cones, but variable and not limited to by its nature subjective opinion of one observer is
this species, straight or curved down towards apex, here not considered to be a valid taxonomic char-
(1–)2.5–4(–5) cm long, with a short, twisted petiole acter. Cephalotaxus koreana was said by its author
and cuneate base, linear-lanceolate, 2.5–4 mm wide, to occur in Korea as well as in Japan, where C. har-
gradually or more abruptly tapering to an acuminate ringtonii can be a shrub as well as a small tree.
or cuspidate apex, coriaceous and more or less rigid, Probably the sugar content of the aril varies among
green above, with two pale green bands below. Mid- individuals or (sub)populations of this species.
rib on the adaxial (upper) side prominent, 0.5 mm
wide, obtusely raised and continuous from base to
Distribution
apex, on the abaxial side nearly flat, 0.5–0.7 mm
wide and continuous. Stomata in two broad bands North and South Korea, Japan, Taiwan.
on the abaxial side, each with 10–15 intermittent TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-NK
white lines, separated by the midrib and bordered KOR-SK TAI
by flat leaf margins. Pollen cones situated in rows of
capitula on the underside of lateral foliage branch-
Ecology
lets; each capitulum on a 3–7 mm long, scaly pedun-
cle inserted in the upper axil of a leaf, subtended by Cephalotaxus harringtonii in its tree form is a com-
ovate, incurved bracts with entire or erose, more or ponent of both broad-leaved (angiosperm) forest
less hyaline margins, bearing 6–12 small, sessile or and coniferous forest, or mixed forest, occurring
short pedunculate, globose cones 3–4 mm diam. in the understorey. Cephalotaxus harringtonii var.
 nana is a spreading shrub on seaside cliffs as well Cephalotaxus harringtonii (Knight ex J. Forbes)
as in mountains over rocky terrain; var. harringtonii K. Koch var. harringtonii. Taxus harringtonii
can also occur as an upright shrub in dense thick- Knight ex J. Forbes, Pinetum Woburn.: 217, t. 63.
ets. Only var. wilsoniana in Taiwan is invariably a 1839, [“harringtonia”]; Cephalotaxus drupacea
small tree in mixed montane forests. It appears that Siebold & Zucc. var. harringtonii (Knight ex
tree forms of this species are increasingly common J. Forbes) Pilg., in Engler, Pflanzenr. IV.5 [18]:
in more southern regions where the winters are 102. 1903. Type: Illustration in J. Forbes, Pinetum
milder. The altitudinal range of var. harringtonii is Woburnense, t. 66. 1839 (holotype). Fig. 83, 84
not well documented; one herbarium collection was
made at 600 m a.s.l. Variety nana occurs from 10 m Cephalotaxus drupacea Siebold & Zucc., Abh. Math.-
266 near the coast to 1900 m a.s.l. in the mountains. In Phys. Cl. Königl. Bayer. Akad. Wiss. 4 (3): 234. 1846.
Taiwan, var. wilsoniana is recorded from 1800 m to Cephalotaxus koreana Nakai, Bot. Mag. (Tokyo) 44:
2700 m a.s.l., so this is a high montane forest tree. 508. 1930.
In Japan, var. harringtonii is common in the under-
growth of forests dominated by Abies sachalinensis
Description
and in open moorland in colder, northern regions.
In S Japan var. harringtonii occurs in mixed forests Shrub to small tree 10 m tall. Leaves often in an up-
with Acer, Quercus, Tsuga, Chamaecyparis, Abies, right V-formation, especially on shoots with pol-
and sometimes, also as a smaller subcanopy tree, len cones, (2.5–)3–4.5(–5) cm long, 3–4 mm wide,
Podocarpus macrophyllus. gradually or sometimes more abruptly tapering to
an acuminate or cuspidate apex.
Uses
Distribution
In Japan, the wood of Harrington’s plum yew is of
minor commercial importance and traditionally North and South Korea; Japan.
used for tool handles and household utensils. The TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-NK
seed arils contain oil which was formerly pressed KOR-SK
from them and used in lamps. This species has long
been cultivated for gardens in Japan, where two of
Conservation
the three varieties here recognized are native. It was
sent to the Leiden Botanical Garden by Philipp von IUCN: LC
Siebold in 1829 as C. drupacea and soon distributed
from there by cuttings, arriving in Ghent, Belgium Cephalotaxus harringtonii (Knight ex J. Forbes)
in 1830. The type of the species C. harringtonii is K. Koch var. nana (Nakai) Rehd., J. Arnold Arbor.
(was) a plant cultivated in the gardens of the Duke 22: 571. 1941. Cephalotaxus nana Nakai, Bot. Mag.
of Bedford at Woburn Abbey, Bedfordshire, Eng- (Tokyo) 33: 193. 1919. Type not designated.
land and represented by plate 66 in the book Pin-
etum Woburnense (Forbes, 1839). It was obtained
Description
from ­Japan by the nursery of Knight & Perry,
Chelsea, London, most likely from seeds obtained Shrubs with upright, suckering stems and layering
from plants in cultivation. It has now been spread lower branches; leaves more or less horizontally ar-
throughout the northern hemisphere and beyond ranged, 1–3(–3.5) cm long, 2–3.5 mm wide, abruptly
and is quite commonly seen in gardens and parks, cuspidate.
including its var. nana and several cultivars. It is
suitable for hedges, especially the smaller var. nana,
Distribution
which spreads readily by suckers.
Japan: Honshu, Hokkaido.
3 varieties are recognized: TDWG codes: 38 JAP-HK JAP-HN
 (ca. 1500–2000 m). These forests are dominated by
Conservation
Fagaceae, Lauraceae, Pinaceae (Pinus, Pseudotsuga,
IUCN: LC Tsuga), and a variety of other trees; C. harringtonii
var. wilsoniana forms a small understorey tree tol-
erant of shade. In more open situations, e.g. after
Uses
forest disturbance, it can hold its own and resprout
This variety is well represented in cultivation, both from base or roots to form a dense bush.
in Japan and abroad, especially in the USA and Eu-
rope. It retains the layering habit in cultivation and
Conservation
therefore will spread out widely if not checked.
The native forests, in which C. harringtonii var. 267
Cephalotaxus harringtonii (Knight ex J. Forbes) wilsoniana is a rare tree, have been under pressure of
K. Koch var. wilsoniana (Hayata) Kitam., Acta conversion after logging to plantations with mainly
Phytotax. Geobot. 26 (1–2): 9. 1974. Cephalotaxus Cryptomeria japonica. The species is known from
wilsoniana Hayata, Icon. Pl. Formos. 4: 22. 1914; several localities concentrated, but not restricted,
Cephalotaxus sinensis (Rehd. & E. H. Wilson) to the northern part of Taiwan, where such conver-
H. L. Li var. wilsoniana (Hayata) L. K. Fu & Nan sions have been widespread. It occurs in several re-
Li, Novon 7 (3): 263. 1997. Type: Taiwan: Nantou, serves, among which is Taroko National Park.
Chia-i Pref., A-li Shan [“Mt. Arisan”], K. Uyematsu IUCN: EN (C2a)
18 (holotype TI).
Uses
Description
This endemic variety from Taiwan is rare in cul-
Small trees to 10 m tall; branches drooping or some- tivation and probably restricted to a few botanic
times pendulous. Leaves mostly spreading horizon- ­gardens.
tally, (1.5–)3–5 cm long, 3–4 mm wide, straight or
slightly falcate, gradually tapering to an acuminate
apex. Cephalotaxus lanceolata K. M. Feng, Acta
Phytotax. Sin. 13 (4): 86. 1975. Cephalotaxus fortunei
Hook. var. lanceolata (K. M. Feng) Silba, Phytologia
Taxonomic notes
68: 27. 1990. Type: China, NW Yunnan, Gongshan,
In the second edition of Flora of Taiwan 1 (1994), as upper Dulongjiang (river), 1900 m a.s.l., Chinese
well as in a horticultural compilation (Tripp, 1995) collector 24347 (holotype PE).
this taxon was treated at species rank. In Flora of
China Vol. 4 (1999) it is treated as a variety under
Etymology
C. sinensis. The morphology of these species is rath-
er similar, but evidence from DNA appears to point The species epithet refers to the shape of the leaves,
to a closer affinity with the Japanese species C. har- resembling the point of a lance.
ringtonii. This classification, first proposed by the
Japanese botanist Siro Kitamura, is here accepted as
Vernacular names
the most likely true reflection of relationships.
Gongshan plum yew; gong shan san jian shan (Chi-
nese)
Distribution
Taiwan.
Description
TDWG codes: 38 TAI
Trees to 20 m tall; trunk to 40 cm d.b.h. Bark thin,
smooth, exfoliating in thin flakes, purplish weath-
Ecology
ering grey-brown. Branches spreading, drooping
Cephalotaxus harringtonii var. wilsoniana occurs or pendulous, forming an open or rounded crown.
in deciduous/evergreen forest at middle elevation Foliage branchlets slender, with grooves between
 decurrent leaf bases, green turning reddish brown.
Conservation
Leaves pectinately arranged in two rows, separated
from each other, forming a V-shape, spreading at Forests have been exploited for timber in the area
45–70° from shoot axis, 4.5–10 cm long, narrowly where this species occurs, but it is unknown to what
lanceolate to linear-lanceolate, straight or slightly extend it has declined as a result. It should eas-
falcate, 4–7 mm wide, very short petiolate, obtuse ily meet the D criterion for Vulnerable (fewer than
at base, very gradually tapering from shortly above 1000 mature individuals) and could well be EN or
base to an acuminate or cuspidate apex, leaf texture even CR under the B criterion if its exact limits of
thin coriaceous, dark green above, pale green with distribution were better known. It is, however, still
two whitish bands below. Midrib obtusely raised problematic to undertake botanic expeditions ac-
268 and continuous on the adaxial (upper) side, 0.6 mm cross the Chinese border into Myanmar.
wide, flat and slightly wider on the abaxial side. Sto- IUCN: VU (D2)
mata in two broad bands of 15–20 intermittent white
lines separated by the abaxial green midrib and bor-
Uses
dered by slightly revolute, green leaf margins. Pol-
len cones unknown. Seed cones solitary at base of No specific uses have been recorded, but due to its
lateral foliage branchlets, on 15–20 mm long, slender potential size as a tree to 20 m tall and 40 cm d.b.h.
peduncles. Bracts several per cone, reduced, 1–1.5 it must have been logged with the rest for timber.
mm long; fertilized ovules surrounded by a green
aril, enclosing the ripening seed, becoming 35–45 ×
20–25 mm, turning soft, greenish brown and indis- Cephalotaxus latifolia L. K. Fu & R. R. Mill,
tinctly striated longitudinally, with a short mucro- Novon 9 (2): 185. 1999. Type: China: Chongking
nate apex. Seeds obovoid or ellipsoid, 30–40 × 16–20 Municipality, Nanchuan Xian, Jinfo Shan,
mm. X. Q. Chen & K. Y. Liang 2463 (holotype PE).

Cephalotaxus sinensis (Rehd. & E. H. Wilson)

Taxonomic notes
H. L. Li var. latifolia W. C. Cheng & L. K. Fu, Acta
The reduction of this species to a variety of C. for- Phytotax. Sin. 13 (4): 86. 1975.
tunei by Silba (op. cit.) is unwarranted; the charac-
ters compared between the two species as given in
Etymology
Flora of China 4: 85 (1999) are distinct. There seem
to be few herbarium specimens; including the type The species epithet describes the broad leaves of this
these are all in China and not easily accessed short species.
of a visit. The Flora Reipublicae Popularis Sinicae
(FRPS) 7: 425 (Cheng & Fu, 1978) gives a line draw-
Vernacular names
ing with foliage and a seed; this illustration was re-
produced in the recently issued illustrations volume Broad-leaved plum yew; kuan ye cu fei (Chinese)
accompanying Flora of China 4.
Description
Distribution
Shrubs to small trees 0.5–6 m tall; trunk d.b.h. to
China: NW Yunnan (Gongshan Drungzu Nuzu 20 cm. Bark thin, exfoliating in narrow strips,
Zizhixian, banks of the upper Dulongjiang); N My- grey-brown. Branches spreading or ascending; fo-
anmar [Burma]. liage branchlets to ca. 10 cm long, stout, glabrous,
TDWG codes: 36 CHC-YN 41 MYA grooved between decurrent leaf bases, green turn-
ing orange-brown. Leaves more or less disposed
in two ranks, spreading horizontally at 65–80° to
Ecology
shoot axis, (1.5–)2–4(–5) cm long, sessile or with a
This species occurs scattered in evergreen broad- very short, twisted petiole and cuneate base, linear
leaved forest at ca. 1900 m elevation. Little else is to linear-lanceolate, 3–4.5(–5.5) mm wide, gradu-
know about its ecology. ally or more abruptly tapering to an acuminate or
­ ucronate apex, ­coriaceous and rigid, dark olive
m
Ecology
green above, with two whitish bands below. Mid-
rib on the adaxial (upper) side prominent, 0.4 mm Cephalotaxus latifolia is a species occurring in
wide, lying in a channel and continuous from base mountainous areas at altitudes between 900 m
to apex, on the abaxial side raised, 0.5–0.6 mm wide and 2400 m a.s.l. It grows in secondary vegetation
and continuous. Stomata in two broad bands on (“thickets” according to Flora of China 4: 86, 1999)
the abaxial side, each with 10–15 intermittent white usually forming a shrub. It may well be more com-
lines, separated by the midrib and bordered by flat mon than previously thought; recent inventories in
or slightly revolute leaf margins. Pollen cones situ- the new Chongqing Municipality (an administrative
ated in rows of capitula on the underside of lateral split from Sichuan Province) have found it there in
foliage branchlets; each capitulum on a 1.5–3 mm several new locations. 269
long, scaly peduncle inserted in the upper axil of a
leaf, subtended by ovate bracts with entire or erose,
Conservation
more or less hyaline margins and a mucronate apex,
bearing 6–10 small, sessile or short pedunculate, This is one of the few conifer species yet to be as-
globose cones 3–4 mm diam. Microsporophylls 5–15 sessed for the IUCN Red List. It seems to be wide-
per cone, each with 2–3 globose, cream coloured spread in S Central and S China but less common
pollen sacs. Seed cones borne 2–6 together at base of than C. fortunei, C. oliveri, and C. sinensis. However,
lateral foliage branchlets, on 2–5 mm long peduncles taxonomic recognition may also play a role; this
with small scale leaves. Bracts several per cone, re- species has formerly been confused with C. har-
duced, 1–1.5 mm long; fertilized ovules surrounded ringtonii, which does not occur in China, or with
by an obovoid, green or purplish aril, enclosing the C. sinensis, and only more recent revision of in par-
ripening seed, becoming 17–20 × 12–15 mm, turning ticular the Chinese herbarium collections has begun
soft, orange-red to purple and smooth or striated, to sort out the correct identity of specimens and
with a short mucronate apex. Seeds (ob)ovoid to therefore of localities.
sub­globose, 12–16 × 9–12 mm, with a small mucro. IUCN: NE

Taxonomic notes Uses

The taxonomic status of this species has been disput- No uses have been recorded of this species.
ed by some authors, who united it with C. sinensis,
to which it bears less resemblance than to C. har-
ringtonii. The latter species commonly has leaves Cephalotaxus mannii Hook. f., Hooker’s Icon.
terminating in a relatively long cusp (as in C. sinen- Pl. 16: t. 1523. 1886. Type: India: Meghalaya, Khasi
sis), while leaves of C. latifolia have an abruptly nar- Hills, Lankhla Woods, G. Mann s.n. (holotype K).
rowed apex with a short cusp being less than 0.5 mm Fig. 85
long. However, variants tending towards either form
of leaf apex occur in C. harringtonii as well as C. lati- Cephalotaxus griffithii Hook. f., Fl. Brit. India 5: 648.
folia, making the distinction less reliable. A com- 1888. [Icon. Pl., ser. 3, 10 (2): t. 1933. 1890].
prehensive revision that would include molecular
evidence is much needed. This investigation should,
Etymology
where possible, refrain from sampling among culti-
vated plants, as their identity or provenance can be This species was named after George Mann, who
muddled to start with. collected the type specimen.

Distribution Vernacular names

China: Chongqing, NW Fujian, N Guangdong, NE Mann’s plum yew; hai nan cu fei (Chinese), Dinh
Guangxi, SE Guizhou, SW Hubei, W Jiangxi. tung, Phi ba mui (Vietnamese)
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHS-
GD CHS-GX CHS-JX
 Description Distribution
Trees to 30(–50?) m tall; trunk to 70(–120) cm d.b.h. China: Guangdong (Xingyi), Guangxi, Yunnan, SE
Bark thin, exfoliating in small or large flakes and Xizang [Tibet]; N Myanmar [Burma]; NE India:
strips, light brown to reddish brown, weathering Arunachal Pradesh (Khasi, Jaintia, Mishmi Hills
grey. Branches spreading or ascending, forming a and Naga Hills), Assam (Manipur, Nagaland); N
rounded or narrow crown. Foliage branchlets up to Thailand; Vietnam; Laos.
25 cm long, slender, grooved between decurrent leaf TDWG codes: 36 CHC-YN CHS-GD CHS-GX CHT
bases, green turning orange-brown. Leaves arranged 40 ASS-MA ASS-NA EHM-AP 41 LAO MYA THA VIE
in two pectinate rows, spreading horizontally and at
270 70–90° to shoot axis, subopposite, (1.5–)2.5–5(–6)
Ecology
cm long, straight or commonly slightly falcate, (2–)
2.5–4 mm wide, flat, base very short petiolate or ses- This widespread species occurs in mixed evergreen
sile, broadly obtuse to nearly truncate, leaf blade or deciduous forests, often in ravines. Its altitudinal
widest below the middle and gradually tapering to range is from 500 m to 2000 m a.s.l. In China trees
a cuspidate apex. Midrib thin but prominent and are described as not reaching taller than 20 m (Flora
continuous from base to apex on the adaxial (upper) of China 4: 87, 1999), but in Thailand trees up to 50
side, 0.4 mm wide, conspicuous but more flattened m tall have been reported. That maximum figure is
on the abaxial side; leaf colour dark green or olive- certainly in need of verification, but trees to 30 m tall
green above, two white or glaucous white bands commonly occur in the undisturbed evergreen sub-
below. Stomata in two broad bands of ca. 20–25 in- montane rainforests of Thailand and Vietnam. Ce-
termittent, white lines, separated by the green mid- phalotaxus mannii can occur on both silicate rocks
rib and bordered by slightly revolute leaf margins. and limestone. It is often associated with Nageia wal-
Pollen cones situated in rows of capitula on the un- lichiana, Taxus wallichiana, Dacrycarpus imbricatus
derside of lateral foliage branchlets; each capitulum and Podocarpus neriifolius on soils derived from the
on a (1–)3–5 mm long, scaly peduncle inserted in the silicate rocks, and with Pseudotsuga sinensis, Nageia
upper axil of a leaf, subtended by ovate, incurved fleuryi, Pinus kwangtungensis, Podocarpus pilgeri,
bracts with entire or erose, more or less hyaline Taxus chinensis, Fokienia hodginsii, and Amento-
margins, bearing 6–8 small, sessile, globose, cream taxus spp. on karst limestone in N Vietnam and S
or light yellow cones up to 4 mm diam. Microsporo- China. In both types of habitat angiosperms play an
phylls 6–12 per cone, each with 3–4 globose, cream important role with numerous tree and shrub spe-
coloured pollen sacs. Seed cones solitary or borne cies as well as epiphytes.
2–3 together at base of lateral foliage branchlets, on
4–10 mm long peduncles. Bracts few per cone, re-
Conservation
duced, 1–1.5 mm long; fertilized ovules surrounded
by an ellipsoid, green aril, enclosing the ripening Globally this species is considered Vulnerable; in
seed, becoming 20–30 × 10–15 mm, turning soft, red China it is considered Endangered, due to the range-
with longitudinal striation. Seeds (ob)ovoid to ellip- wide reduction of forests. Throughout its range,
soid, sometimes laterally compressed, 18–28 × 8–14 C. mannii is restricted to small populations in which
mm, with a mucronate or cuspidate apex. the largest trees (to 30 m tall and over 100 cm d.b.h.)
are often targeted for their timber. Stripping of bark
is often fatal to the trees and this type of harvest, like
Taxonomic notes
logging due to slow growth, is unsustainable. The
This species has sometimes been treated in a wid- principal threat, however, is conversion of habitat
er sense to include C. hainanensis, e.g. in Flora of to agriculture and resulting severe forest fragmen-
China 4 (1999), but this taxon is here treated as a tation. There are not enough forest reserves of suf-
distinct species. Cephalotaxus is a difficult genus ficient size and integrity to safeguard this species
morphologically and a comprehensive critical revi- at present (Fu & Jin, 1992; Nguyen Tien Hiep et al.,
sion including DNA-based analyses is long overdue. 2004). The current global assessment dates from the
1990’s and a renewed assessment under the IUCN slightly rugose, stiff; leaf colour dull green above,
2001 criteria is very urgent. It is likely that habitat pale green below with two whitish bands. Midrib
reduction has accelerated and that the species would on the adaxial (upper) side narrow (less than 0.4
move into a higher category of threat. mm wide), raised but fading towards apex, on the
IUCN: VU (A1d) abaxial side 0.7–0.8 mm wide, flat. Stomata in two
broad bands of 13–17 intermittent lines separated by
the abaxial midrib and two 0.7 mm wide margins
Uses
and of equal width to these. Pollen cones situated
This conifer produces high quality insect resistant in rows of capitula on the underside of lateral foli-
timber which is used for quality furniture, fine crafts age branchlets; each capitulum on a 1–3 mm long,
and tool handles. The seeds have medicinal qualities scaly peduncle inserted in the upper axil of a leaf, 271
and in Hainan the bark is used to treat fever. The subtended by ovate, incurved bracts with entire or
species is eminently suitable for use in horticulture, erose, more or less hyaline margins, bearing 3–6
but is seldom seen in cultivation outside Asia. Some small, sessile, globose, pink or light brown cones
plants may be grown under its taxonomic synonym, up to 2 mm diam. Microsporophylls 4–9 per cone,
C. griffithii Hook. f., which was originally described each with 3(–4) globose, pink coloured pollen sacs.
from the Mishmi Hills in Assam, India. At the Royal Seed cones solitary or in groups at base of lateral fo-
Botanic Garden, Edinburgh cuttings are being prop- liage branchlets, on 3–7 mm long, curved peduncles.
agated from sources in Vietnam. Bracts several per cone, reduced, 1–1.5 mm long;
fertilized ovules surrounded by an obovoid, green
or grayish aril, enclosing the ripening seed, becom-
Cephalotaxus oliveri Mast., Bull. Herb. Boiss. 6: ing 22–27 × 14–18 mm, turning soft, orange-red to
270. 1898. Type: China: Hubei, Changyang T. Z., red with longitudinal striation. Seeds (ob)ovoid to
A. Henry 7479 (lectotype K, designated here). ovoid, sometimes subglobose, 18–23 × 10–15 mm,
with a mucronate apex.
Etymology
Taxonomic notes
The species epithet commemorates F. W. Oliver, a
botanist who collected in China. Records of this species from Vietnam are referable
to C. mannii.
Vernacular names
Distribution
Oliver’s plum yew; bi zi san jian shan (Chinese)
China: Chongqing, N Guangdong, Guizhou, W Hu­
bei, Hunan, E Jiangxi, S and W Sichuan (especially
Description
Emei Shan), E Yunnan.
Shrubs or small trees to 4 m tall. Bark thin, becom- TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
ing scaly with thin flakes, yellow weathering greyish SC CHC-YN CHS-GD CHS-HN CHS-JX
brown. Branches spreading to ascending, forming
a bushy crown. Foliage branchlets slender, lateral
Ecology
branchlets (sub)opposite, often in a plane, up to 15
cm long but usually shorter than 10 cm, grooved Cephalotaxus oliveri occurs in evergreen and de-
between decurrent leaf bases, yellowish green turn- ciduous broad-leaved forests, coniferous forests,
ing yellow to light brown. Leaves in two opposite, and mixed forests, at altitudes between 300 m and
distichous ranks at 60–80° to shoot axis, mostly of 1800 m a.s.l. As it apparently never grows taller
equal length and touching each other from base to than about 4 m, it is an understorey shrub, occur-
near apex, (1.5–)2–3(–3.5) cm long, (2.3–)2.7–3.5(–4) ring in the shrub layer mixed with several genera
mm wide, straight or slightly falcate and convex, of angiosperms such as Rhododendron, Camellia,
sessile or nearly so, base truncate or weakly cordate, Cotoneaster, Deutzia, Lonicera, Berberis, Buddleia,
flat margins parallel to near the short, minutely cus- Euonymus, Hydrangea, Prunus and many other spe-
pidate or mucronate apex, leaf texture coriaceous, cies.
 r­ eddish to grey-brown. Branches spreading, droop-
Conservation
ing or ascending; foliage branchlets short or long and
This species has a relatively wide geographical range slender, glabrous, grooved between decurrent leaf
and broad ecological amplitude, yet it has suffered bases, green turning yellowish brown. Leaves more
from widespread deforestation as well as direct ex- or less disposed in two ranks, sometimes upright
ploitation for its bark. The IUCN criteria used are forming a V-formation especially on branchlets with
those of 1994; at those of 2001 a 50% or greater re- male cones, straight or slightly falcate, (1–)1.8–5(–7)
duction in three generations has to be inferred or cm long, with a short, twisted petiole and cuneate
suspected in order for the species to be listed as VU, base, linear-lanceolate or linear, 2–4 mm wide, more
and it remains to be seen if this can be substantiated or less abruptly tapering to a long acuminate or
272 in the next assessment. cuspidate apex, coriaceous but relatively soft, green
IUCN: VU (A1c-d) above, with two white or rarely pale green bands be-
low. Midrib on the adaxial (upper) side prominent,
0.4 mm wide, obtusely raised and continuous from
Uses
base to apex, on the abaxial side nearly flat, 0.5–0.6
This shrubby species is being exploited for its bark, mm wide and continuous. Stomata in two broad
which contains anti-carcinogenic alkaloids, for me- bands on the abaxial side, each with 12–15 intermit-
dicinal purposes; it is also in cultivation as an or- tent white lines, separated by the midrib and bor-
namental shrub both in and outside China. It was dered by flat or slightly revolute leaf margins. Pollen
introduced to Britain and the USA by E. H. Wilson cones situated in rows of capitula on the underside
but apparently has not been successful, probably due of lateral foliage branchlets; each capitulum on a 1–3
to cold winters. It should certainly be suitable as a mm long, scaly peduncle inserted in the upper axil
garden shrub in countries with mild and (nearly) of a leaf, subtended by ovate, incurved bracts with
frost-free winters. entire or erose, more or less hyaline margins, bear-
ing 6–7 small, sessile, globose, pink or light brown
cones up to 3 mm diam. Microsporophylls 4–11 per
Cephalotaxus sinensis (Rehd. & E. H. Wilson) cone, each with 2–3(–4) globose, pink coloured pol-
H. L. Li, Lloydia 16 (3): 162. 1953. Cephalotaxus len sacs. Seed cones solitary or borne 2–6 together
drupacea Siebold & Zucc. var. sinensis Rehd. & at base of lateral foliage branchlets, on 3–8 mm long,
E. H. Wilson, in Sargent, Pl. Wilson. 2: 3. 1914; curved peduncles. Bracts several per cone, reduced,
Cephalotaxus harringtonii (Knight ex J. Forbes) 1–1.5 mm long; fertilized ovules surrounded by an
K. Koch var. sinensis (Rehd. & E. H. Wilson) ellipsoid, green or grayish aril, enclosing the ripen-
Rehd., J. Arnold Arbor. 22: 571. 1941. Type: China: ing seed, becoming 18–25 × 9–16 mm, turning soft,
W Sichuan, Baoxing, [“Mupin”], E. H. Wilson 1115 red or purple with 6 longitudinal ridges. Seeds (ob)
(holotype A). ovoid to ellipsoid, rarely subglobose, 16–22 × 8–14
mm, with a mucronate or cuspidate apex.
Etymology
Distribution
The species epithet refers to China, its native coun-
try. S Central, S (including Hainan), SW, and SE China.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
SC CHC-YN CHS-AH CHS-FU-CHS-GD CHS-GX CHS-
Vernacular names
HE CHS-HK CHS-HN CHS-JS CHS-JX CHS-ZJ
Chinese plum yew; cu fei (Chinese)
Ecology
Description
Cephalotaxus sinensis is very widespread in SE Chi-
Shrubs to small trees 1.5–15 m tall; trunk d.b.h. of na and occurs in a range of habitats between 200 m
monopodial trees to 120 cm. Habit of shrubs erect a.s.l. on the eastern coast to 2800(–3200) m a.s.l. in
and spreading; trees with a wide, open and round- the mountains of Sichuan and Yunnan. It is common
ed crown. Bark thin, exfoliating in narrow strips, in moist woodlands along streams at lower altitudes,
as well as in montane coniferous or mixed forests l­ocal trade, largely because sizable trees are very scat-
and in shrubby thickets on mountain slopes, on tered. The wood is used to make furniture, farm tools
granite, sandstone or limestone. In Zhejiang Prov- such as handles, other utensils and crafts like wood
ince, in the far east of its range, it is a minor compo- turning. Similarly to the other species, the seed arils
nent in diverse mixed mesophytic forest below 750 yield oil that was traditionally burnt in lamps. Alka-
m a.s.l. with e.g. Acer, Carya, Celtis, and Quercus s. loids in the foliage, roots and seeds are extracted for
l. (including Castanopsis) as well as the rare conifer medicinal purposes, e.g. as treatment of leukemia.
Pseudolarix amabilis and, in the Tienmu Shan, per- This species is in cultivation in China on a limited
haps the last wild-growing Ginkgo biloba trees. scale, mainly in botanic gardens. Outside China it
is rarely cultivated, but plants belonging to this spe-
cies may have been confused with C. harringtonii. 273
Conservation
Its long acuminate leaf tips, however, should distin-
IUCN: LC guish even sterile plants from C. harringtonii. Being
often from more southerly regions it is not as hardy,
but this may depend on provenance and some of the
Uses
high altitude populations of Sichuan and Yunnan
Given the reported size of the diameter (d.b.h.) of can be grown as far north in the USA as Massachus-
some tree trunks, it is obvious that its wood is put setts, where Ernest Wilson first introduced the spe-
to use, but it is not commercially important beyond cies as C. drupacea var. sinensis.
 Chamaecyparis Spach, Hist. Nat. Vég. Phan. 11: 329. 1841. Type: Chamaecyparis thy­
oides (L.) Britton, Sterns & Poggenb. (Cupressus thyoides L.) [Chamaecyparis sphae­
roidea (Spreng.) Spach (nom. illeg.) (Thuja sphaeroidea Spreng.)] (Cupressaceae).

Retinispora Siebold & Zucc., Fl. Japon. 2 (5): 36. 1844. 1b. Seed cone scales 8–14(–16) in number, mature
Type: Retinispora obtusa Siebold & Zucc. [Chamae- cones 7–14 mm diam 3
cyparis obtusa (Siebold & Zucc.) Endl.] 2a. Smallest branchlets plagiotropic (in flat sprays);
ultimate branchlets often unilateral C. pisifera
Greek: chamae- = lowly, creeping; cyparis = cypress. 2b. Smallets branchlets irregularly disposed; ulti-
274 mate branchlets alternating C. thyoides
3a. Seed cones widely open, with gaps between
Description
scales as wide as or wider than scales. Leaf
Evergreen, monoecious trees; trunk monopodial. gland present; leaf apices of laterals more or less
Resin cavities in leaves. Bark fissured, fibrous or acute C. lawsoniana
scaly, exfoliating in long strips or flakes, reddish 3b. Seed cones not widely open, with gaps between
brown. Branches slender, persistent, forming a py- scales narrower than scales. Leaf gland absent
ramidal, conical or rounded crown (Massart’s mod- or very obscure; leaf apices of laterals obtuse 4
el). Fastigiate forms restricted to cultivation. Foliage 4a. Seed cones longer than wide when closed; seed
branches plagiotropic, drooping or pendulous, (sub) wings nearly equal C. formosensis
ultimate branchlets flattened, covered with scale 4b. Seed cones always globose; seed wings often
leaves. Leaves decussate, imbricate, dimorphic, with unequal C. obtusa
facials smaller than laterals, appressed, with apices
of laterals spreading above apices of facials; mar-
gins entire; stomata inconspicuous, mostly on the Chamaecyparis formosensis Matsum., Bot. Mag.
underside of the (leaf-covered) branchlets. Pollen (Tokyo) 15: 137. 1901. Retinispora formosensis
cones terminal, solitary, very small, short cylindri- (Matsum.) A. V. Bobrov & Melikyan, Komarovia 4:
cal; micro­sporophylls 8–16, decussate, bearing 2–3 74. 2006. Type: Taiwan: Nantou, Chia-i Pref., Yu-
abaxial, yellow or red pollen sacs. Seed cones termi- Shan, [“Mt. Morrison”], S. Honda s.n. (lectotype
nal, solitary, globose to ellipsoid-ovoid, small, with TI). Fig. 86, 87
peltate, parting scales. Bract-scale complexes (6–)
8–12(–16), decussate, peltate, with a quadrangular,
Etymology
rhombic or polygonal outline, abaxially depressed
with a small central umbo (bract tip), adaxially with The species epithet indicates its origin as from For-
faint seed marks near base; the ultimate pair sterile. mosa, an earlier name for Taiwan; perhaps with a
Seeds moderately numerous, with 2 lateral, narrow pun to Latin: formosus = handsome or well formed.
wings. Seedlings with 2 cotyledons.
Vernacular names
5 species.
Taiwan Cypress, Formosan Cypress; Beniki; hong
gui (Chinese)
Distribution
North America: (disjunct) E North America; in
Description
W North America in Oregon and California. Asia:
Japan, Taiwan. Trees to 55–60 m tall; trunk monopodial, branching
low, occasionally forked, up to 6–7 m d.b.h. in a few
very old trees. Bark soon flaky, on large trees more
Key to the species of Chamaecyparis
than 20 cm thick towards base of trunk, fissured, fi-
1a. Seed cone scales 4–8(–10) in number, mature brous, exfoliating in long strips, light reddish brown
cones 4–8 mm diam 2 weathering grey. Branches spreading or curved
down, higher order branches drooping, forming a
dense pyramidal crown in young trees, eventually the seed wings are more developed and presumably
becoming broadly conical or sympodial and domed more effective in dispersal as a result. It is the largest
in very old specimens. Foliage branches numerous, conifer (measuring stem diameter) growing in Asia
spreading and drooping; plagiotropic branchlets al- and possibly the longest-lived as well. A detailed
ternating, smallest ones often unilateral on second study of its biology is very desirable.
highest order, gradually shortening forming taper-
ing planate sprays, covered with green leaves; ulti-
Distribution
mate lateral branchlets partly deciduous after 4–6
years. Leaves decussate, imbricate, decurrent, scale- Taiwan: central mountains.
like, 1–3 × 0.5–1 mm on ultimate branchlets, up to TDWG codes: 38 TAI
10 mm long on leading shoots, dimorphic; facials 275
smaller than laterals, rhombic to lanceolate, carinate,
Ecology
obtuse-acuminate, appressed or with a free apex,
with an inconspicous abaxial gland; laterals connate In mixed coniferous forest of the cool temperate
proximally, spreading above the apex of facials, con- coniferous forest belt between 1700 m and 2900 m
duplicate, lanceolate, recurved and incurved at the a.s.l., with Chamaecyparis obtusa var. formosana and
mostly free apex, eglandular or obscurely glandular; C. formosensis as the dominant species, associated
margins entire; leaves amphistomatic, stomata in- with Calocedrus formosana, Cunninghamia konishii,
conspicuous except for a few exposed lines on un- and Taiwania cryptomerioides, in the upper part of
derside of facials; leaf colour light grey-green or dull the zone also Tsuga chinensis; in the understorey
green. Pollen cones terminal, solitary, ovoid-oblong, occur Photinia davidiana var. niitakayamensis and
2–3 × 1–1.5 mm, yellowish green turning brown; the bamboo Yushania niitakayamensis, but often lit-
microsporophylls 8–12, decussate, peltate, sub­ tle else. Chamaecyparis formosensis is exceptionally
orbicular, with minutely denticulate margins, with long-lived, estimated in excess of 3000 years, and
3 abaxial yellow pollen sacs on the lower margin. has a regeneration strategy similar to e.g. Sequoia-
Seed cones terminal on branchlets with unmodi- dendron, although the disturbance factors are not
fied leaves, solitary, maturing within one year, cadu- well known. Soils are usually slightly acidic, derived
cous, subglobose to ellipsoid-ovoid, (6–)10–12 × 5–8 from igneous rock or shale. The climate is cool and
mm with opened scales, from purplish ripening to wet, with abundant rainfall throughout the year and
brown. Bract-scale complexes (8–)10–14(–16), dec- occasional snowfall in winter.
ussate, parting and spreading at right angles from
axis when mature, subpeltate to peltate, rhombic in
Conservation
outline, 3–5.5 mm wide; abaxial surface depressed,
with a central recurved umbo (bract tip 0.5–0.7 The number of mature trees in natural forest stands
mm); margin undulating; base conical; adaxial face has been greatly reduced by felling. The species oc-
grooved and striated, lustrous brown, lacking seed curs as scattered individuals in mixed coniferous
marks. Seeds (1–)2(–3) at the basis of each scale forest and has been selectively logged for its desir-
(15–20 develop per cone), slightly flattened, oblong, able wood used in traditional oriental building.
1.5–2.5 mm long, reddish brown, surrounded by 2 Growth is slow, especially in mature to over-mature
lateral, nearly equal thin wings 0.5–1 mm wide. phases of its long life cycle. Several very large, se-
nescent individuals remain as ‘natural monuments’
forming a tourist attraction, but protection of more
Taxonomic notes
old growth forests containing this species is urgently
This species may be more closely related to Chamae- needed. The species is being grown in plantations.
cyparis pisifera of Japan than to its congener on the IUCN: EN (A1c)
island of Taiwan: C. obtusa var. formosana. The two
species in Taiwan, C. formosensis and C. obtusa, are
Uses
sympatric but stay well separate taxonomically. The
much greater size of C. formosensis is related to lon- The wood (timber) of this species is highly prized
gevity and this is probably genetically determined; for traditional oriental buildings such as temples
its seed cones are also larger and more ellipsoid and and shrines. It is uncommon in cultivation outside
 Taiwan, where this species is now being replanted planate sprays, covered with green leaves; ultimate
in attempts to restore it from excessive logging in lateral branchlets partly deciduous after 4–6 years.
the past. As an ornamental it is attractive, but slow Leaves decussate, imbricate, decurrent, scale-like,
growing and similar to Sawara cypress (C. pisifera) 2–3 × 1–1.5 mm on ultimate branchlets, up to 20 mm
from Japan, which is much more common as a tree long on leading shoots, dimorphic; facials smaller
for the garden. To increase its use as an ornamen- than laterals, rhombic to lanceolate, obtuse-acumi-
tal requires marketing efforts and with so many nate, appressed, with a conspicous, oval, yellowish,
cultivars already available from other species in transparant, non-active abaxial gland; the laterals
the genus, this is not an easy task, and one which connate proximally, spreading above apex of facials,
commercial growers will be reluctant to undertake. conduplicate, broadly falcate to lanceolate, recurved
276 Outside Taiwan, fine specimen trees grow at Hillier’s and incurved at the appressed apex, less conspicu-
Arboretum in the south of England, where ground ously glandular; margins entire; leaves amphisto-
frost is quite common; this species is apparently not matic, stomata inconspicuous, concentrated on the
as tender as is commonly believed. partly hidden bases and margins of leaves; leaf col-
our light green or lustrous dark green; margins more
or less glaucous, stomatal zones greenish white. Pol-
Chamaecyparis lawsoniana (A. Murray bis) Parl., len cones very numerous, terminal, solitary, ovoid-
Ann. Mus. Imp. Fis. Firenze 1: 181. 1864. Cupressus oblong, 3–4 × 1.5–2 mm, yellowish green turning
lawsoniana A. Murray bis, Edinburgh New Phi­ purplish black; microsporophylls 12–16, decussate,
los. J., n.s., 1: 292. 1855; Retinispora lawsoniana peltate, with minutely denticulate margins, with 2–3
(A. Murray bis) A. V. Bobrov & Melikyan, Koma­ abaxial large, red pollen sacs on the lower margin.
rovia 4: 74. 2006. Type: Illustration in A. Murray, Seed cones terminal on branchlets with unmodified
Edinburgh New Philos. J. n.s. 1, t. 10, f. 1–4. 1855 leaves, solitary, maturing within one year, caducous,
(lectotype); USA: California, W. Murray s.n. [3] subglobose, (6–)7–11(–14) mm with opened scales,
(epitype E). Fig. 88, 89 from glaucous purplish (bracts) and yellow ripen-
ing to grey-brown. Bract-scale complexes 8–10(–12),
decussate, ultimate pair often sterile and fused,
Etymology
parting and spreading at varying angles from axis
This species was named after Charles Lawson (1794– when mature, subpeltate to peltate, polygonal (4–5
1873), an English nurseryman who grew several ear- angular) in outline, up to 6–7 mm wide, abaxial sur-
ly introductions of conifers. face slightly depressed, rugose, with a central small
umbo (bract tip 0.5–1 mm); base conical; adaxial
surface grooved and striated, pinkish brown, with
Vernacular names
faint seed marks near the base. Seeds 2–4 at the ba-
Port Orford Cedar, Port Orford White-cedar, Or- sis of each scale (15–30 develop per cone), slightly
egon Cedar, Lawson Cypress flattened, ovoid, with acute apex, 2.5–3 × 2 mm long,
yellowish, with 2 oblong, darker resin glands on
each side, surrounded by 2 lateral, nearly equal thin
Description
wings 1–1.5 mm wide.
Trees to 60–70 m tall; trunk monopodial, large
trees can be buttressed at base, up to 4–4.5 m d.b.h.
Distribution
Bark soon flaky, on large trees to 35–40 cm thick
towards base of trunk, deeply fissured, fibrous, ex- USA: SW Oregon, NW California, more extensively
foliating in long strips, dark red-brown weathering in Oregon (Coos Co., Curry Co.), from the Pacific
grey-brown. Branches slender, curved down; higher coast to Mt. Shasta.
order branches drooping or pendulous, forming TDWG codes: 73 ORE 76 CAL
a dense conical crown in young trees, eventually
becoming broadly columnar in large trees. Foliage
Ecology
branches numerous, drooping or pendulous, plagio-
tropic branchlets alternating and smallest ones often This species is usually growing in mixed conifer-
unilateral, gradually shortening forming tapering ous forest of different types, with e.g. Abies concolor,
A. grandis, Picea sitchensis, P. breweriana, Pinus
Uses
spp., Pseudotsuga menziesii, Sequoia sempervirens,
and Tsuga heterophylla, in various mixtures; also in The timber of this species is highly valued in East
mixed conifer/ angiosperm forest with e.g. Quercus Asian countries, especially Japan, for construction
and Lithocarpus densiflorus. Understorey vegetation of traditional buildings. There is a major inter­
is usually made up of the ericaceous shrubs Rho- national trade, especially with Japan, where it fetch-
dodendron and Vaccinium, Rhamnus spp., Rubus es extremely high prices. There is probably no other
spectabilis, Gaultheria shallon and Quercus spp. and conifer species which has given rise to more cultivar
more locally Taxus brevifolia; a herbaceous flora is forms than C. lawsoniana, several hundred of which
also commonly present. Except when growing with are listed in recent compilations (Leslie, 1992; Welch
a dominance of Pinus, the forests are dense. Habitats & Haddow, 1993). These lists were outdated upon 277
less favourable to forest development, e.g. bogs and publication as the number of new cultivar forms
dry sand dunes, can still be occupied by Chamae- arising and sold by nurseries seems to grow expo-
cyparis lawsoniana, which will then grow much nentially. Registration of these by the Conifer Reg-
slower and less tall in the communities that prevail istrar of the Royal Horticultural Society in England,
there. Best development is on wet soil types with the acknowledged authority in the world of conifer
subsurface seepage, where the species can become horticulture, has become an almost impossible task.
dominant; on drier sites the underlying rock is of- Many older cultivars only exist, if at all, as one or a
ten ultramafic. Its altitudinal range is from near sea few surviving individual shrubs or trees in gardens
level to ca. 1500 m a.s.l. The climate is strongly oce- or parks and will become ‘extinct’ if not vegetatively
anic but summers tend to be warm and dry (June- propagated. Numerous names would therefore have
August); total precipitation ranges from 1000–2250 to be deleted from the register if it could be shown
mm annually, at higher altitudes a substantial part that no surviving plants exist. To demonstrate that
is snow. is even more difficult than to list the new cultivars.

Conservation
Chamaecyparis obtusa (Siebold & Zucc.) Endl.,
Over-exploitation of mature and old growth stands, Syn. Conif.: 63. 1847.
exacerbated with a more recent infestation by the
oomycete pathogen Phytophthora lateralis, which
Etymology
was accidentally introduced in 1952, has led to a
steady decline over more than 150 years. Opening up The species epithet refers to the obtuse (blunt) leaf
areas formerly inaccessible with vehicles is part of apices.
the problem, as the pathogen gets transported into
new stands more quickly than would otherwise have
Vernacular names
been possible. For this reason the establishment of
some 12 Research Natural Areas on National Forest Hinoki Cypress; Hinoki (Japanese)
lands has been only partly effective in halting the
decline. An attempt to have this species listed in Ap-
Description
pendix II of CITES by the Oregon Natural Resourc-
es Council in 1994 has so far remained unsuccessful. Trees to 40–50 m tall; trunk monopodial, large trees
Silvicultural practices will urgently have to be re- can be buttressed at base, up to 2 m d.b.h. Bark
vised to make the harvest of this species for timber soon flaky, on large trees fissured, scaly, exfoliating
sustainable. Plantation forestry will largely have to in strips and flakes, red-brown weathering grey-
replace harvesting natural stands despite relatively brown. Branches long, slender, ascending, spread-
slow early growth rates, in order to separate for- ing or curved down in lowest part of crown; higher
estry from areas of infestation and to allow natural order branches drooping or pendulous, forming a
stands to recover. Control of the international trade dense pyramidal crown in young trees, eventually
is ­essential. becoming broadly rounded in large trees. Foliage
IUCN: VU (A1d-e, A2e) branches numerous, drooping or pendulous; plagio-
tropic branchlets alternating and smallest ones ­often
 ­ nilateral, forming irregularly tapering planate
u sites. Both occur in mixed conifer/angioserm for-
sprays, covered with green leaves; ultimate lateral ests, in which the conifers are mostly emergents and
branchlets partly deciduous after 4–6 years. Leaves the angiosperms form lower layers of canopy. More
decussate, imbricate, decurrent, scale-like, coria- specific information is given with the descriptions
ceous, 0.8–3 × 0.5–1.5 mm on ultimate branchlets, up of each variety.
to 10 mm long on leading shoots, dimorphic; facials
smaller than laterals, rhombic to oblong, obtuse,
Uses
appressed, eglandular; laterals connate proximally,
spreading above apex of facials, conduplicate, broad- The wood of Chamaecyparis obtusa has been used
ly falcate to lanceolate, recurved and incurved at the for centuries in construction of temples and other
278 appressed obtuse apex, eglandular; margins entire; traditional buildings because of its fine quality and
leaves amphistomatic, stomata concentrated on the high durability in outdoor conditions. The Japanese
partly hidden bases and margins of leaves; leaf col- have largely turned to sources outside Japan to ob-
our green or grey-green, stomatal zones greenish tain timber of related species, especially those oc-
white on underside of branchlets. Pollen cones ter- curring on the Pacific coast of North America. This
minal, solitary, subglobose, 2–3 mm, orange-brown species is among the most widely used in horticul-
or purplish turning dark brown; microsporophylls ture and numerous cultivars, obtained in Japan,
6–8, decussate, peltate, with entire, rounded mar- Europe, the United States. and New Zealand, are in
gins, with 3 abaxial large, yellow or reddish pollen the trade. The species is particularly suitable for the
sacs on the lower margin. Seed cones terminal on clonal propagation of dwarf forms (‘tennis ball coni-
branchlets with unmodified leaves, solitary, matur- fers’) with compact growth selected from cuttings
ing within one year, caducous, subglobose or glo- from witches brooms; these grow often very slowly
bose, (7–)8–11(–13) mm with opened scales, from and are therefore considered ideal for pot-grown
yellowish green (bracts) and yellow ripening to red- patio plants, rockeries, etc. The number of cultivar
dish brown or dark brown. Bract-scale complexes names under this species is rapidly outgrowing the
8–10, decussate, parting and spreading at varying capacity to list them and even the most comprehen-
angles from axis when mature; distal 2 connate and sive register of cultivated conifers, so far compiled
usually sterile, peltate, polygonal (4–5-angular) in and published by the Royal Horticultural Society
outline, 3–6 mm wide; abaxial surface depressed, in England for the names from letters A-J (Leslie,
rugose, with a central small umbo (bract tip 0.5–1 1992), cannot claim to be nearly complete.
mm); base conical; adaxial surface grooved and stri-
ated, red-brown, with light grey seed marks near 2 varieties are recognized:
base. Seeds 2(–3) at the basis of each scale (15–20 de-
velop per cone), flattened, ovoid, with resin vescicles Chamaecyparis obtusa (Siebold & Zucc.) Endl.
or longitudinal grooves, 2.5–4 × 2 mm long, lustrous var. obtusa. Retinispora obtusa Siebold & Zucc., Fl.
reddish brown, surrounded by 2 lateral, unequal or Japon. 2 (5): 38, t. 121. 1844. Type: Japan: Honshu,
nearly equal thin wings 1–1.5 mm wide. [locality not stated], P. F. von Siebold s.n. (holotype
not located, isotype K). Pl. 11
Distribution
Description
Japan: Honshu, Kyushu; Taiwan.
TDWG codes: 38 JAP-HN JAP-KY TAI Leaves on ultimate branchlets 1.5–3 mm long, 1–1.5
mm wide. Seed cones subglobose, 9–12(–13) mm
diam. when open. Larger seeds are correlated with
Ecology
greater cone size.
The two varieties of this species occur in differ-
ent (micro)habitats: C. obtusa var. obtusa is gener-
Distribution
ally occupying more xeric sites on ridges or slopes;
C. obtusa var. formosana forms extensive forests in Japan: S Honshu, Shikoku, Kyushu.
atmospherically damp and often edaphically wet TDWG codes: 38 JAP-HN JAP-KY JAP-SH
 279
5

2
9
10

Pl ate 11. Chamaecyparis obtusa var. obtusa. 1. Habit of tree. 2. Branch with foliage. 3. Branchlet with
leaves. 4. Juvenile leaves. 5. Adult leaves. 6. Branchlet with leaves and pollen cones. 7. Microsporophyll with
open pollen sacs and pollen. 8. Seed cones. 9. Seed cone scale. 10. Seeds.
 Ecology Distribution
In mixed coniferous and conifer-broad-leaved for- Taiwan.
est with Chamaecyparis pisifera, Pinus densiflora, TDWG codes: 38 TAI
P. parviflora, Abies firma, Pseudotsuga japonica,
Tsuga sieboldii, Sciadopitys verticillata (locally), Aes-
Ecology
culus turbinata, Acanthopanax sciadophylloides, Acer
spp., Fagus japonica, Quercus sp., Stewartia mona- In mixed coniferous or conifer-broad-leaved forest in
delpha, and Magnolia obovata, on Yakushima with the cool temperate coniferous forest belt at altitudes
Cryptomeria japonica. Variety obtusa is widespread from 1800 m to 3000 m a.s.l., with Chamaecyparis
280 on older montane formations, but virtually absent formosensis, Calocedrus formosana, Cunninghamia
from (recent) volcanoes (Wilson, 1916), indicating konishii, Taiwania cryptomerioides, Alnus henryi,
its niche in the late phases of mountain forest suc- Schima superba, Photinia davidiana, Rhododendron
cession, where if undisturbed it becomes a canopy formosanum, Pasania sp., and the alpine bamboo
emergent rising above most other conifers. It occurs Yushania niitakayamensis. Soils are relatively deep
in cool, moist ravines and on slopes with a northerly and well-drained loams or loamy sand. The climate
aspect, as well as on more xeric ridges, in relatively is cool, very moist, with cloud-cover resulting in fog
deep, rich loams over sedimentary or metamorphic and rain much of the year and very high precipita-
or old volcanic rock. The altitudinal range is from tion exceeding 4000 mm per year.
80 m to 2500 m a.s.l.
Conservation
Conservation
Logging has caused a considerable decline in many
IUCN: NT of the more accessible stands. After massive but un-
quantified destruction the total area of occupancy
Chamaecyparis obtusa (Siebold & Zucc.) Endl. var. (AOO, in mixed stands with other conifers) 50 years
formosana (Hayata) Hayata, Fedde’s Repert. Sp. ago was ca. 50,000 ha for C. obtusa var. formosana
Nov. Regni Veg. 8 (23–25): 365. 1910. Chamaecyparis and C. formosensis combined (Lee, 1962). It is almost
obtusa (Siebold & Zucc.) Endl. var. obtusa f. formo­ certainly less than that now, as the decline of natural
sana Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25 (19): forest has continued from 66% to 52% of the land
208. 1908; Chamaecyparis obtusa (Siebold & Zucc.) surface (Yang & Pan, 1996). Old growth forest is be-
Endl. subsp. formosana (Hayata) H. L. Li, Taiwania ing replaced by secondary vegetation; where active
1: 305. 1950. Type: Taiwan: [“Shinko, Shirakku”], reforestation is undertaken the emphasis is usually
T. Kawakami & U. Mori 1329 (lectotype TI). Fig. 86 on faster growing conifers (Cryptomeria, Cunning-
hamia, Pinus). Important stands of Chamaecyparis
Chamaecyparis taiwanensis Masam. & S. Suzuki, spp. are protected in the Yuanyang Lake Reserve.
Sylvia 4: 57, t 1, f. B3–4, t. 2, f. B54. 1933; Retinispora IUCN: VU (A1d)
taiwanensis (Masam. & S. Suzuki) A. V. Bobrov &
Melikyan, Komarovia 4: 74. 2006.
Chamaecyparis pisifera (Siebold & Zucc.) Endl.,
Syn. Conif.: 64. 1847. Retinispora pisifera Siebold &
Vernacular names
Zucc., Fl. Japon. 2 (5): 39, t. 122. 1844. Type: Japan:
Taiwan bian mai (Chinese) Honshu, [loc. unknown], P. F. von Siebold s.n.
(holotype not located, isotype K).
Description
Etymology
Leaves of ultimate branchlets 0.8–1.5 mm long, 0.5–1
mm wide. Seed cones globose, 7–9 mm in diam. The species epithet may refer to “pear-shaped” seeds.
when open. Smaller seeds are correlated to smaller
cone size.
 tip 0.5 mm or invisible); base conical; adaxial ­surface
Vernacular names
striated, light brown, with faint seed marks near
Sawara Cypress; Sawara (Japanese) base. Seeds 1–2 at base of each scale (10–15 develop
per cone), slightly flattened, ovoid, 2 × 1.3 mm; apex
acute; brown, with 3–6 lighter resin vesicles on each
Description
side and a concave hilum at base; wings 2, lateral,
Trees to 40–50 m tall; trunk monopodial, large trees nearly equal thin, translucent, 1.5–2 mm wide.
buttressed at base or not, up to 2 m d.b.h. Bark soon
flaky, on large trees fissured, fibrous, exfoliating in
Distribution
long strips, reddish brown weathering grey-brown.
Branches slender, ascending, spreading or curved Japan: Honshu, Kyushu. 281
down; higher order branches spreading or droop- TDWG codes: 38 JAP-HN JAP-KY
ing, forming a dense conical crown in young trees,
eventually becoming broadly columnar in large
Ecology
trees. Foliage branches numerous, drooping or
pendulous; plagiotropic branchlets alternating and This species has a more scattered distribution in the
smallest ones often unilateral, gradually shortening forests than C. obtusa and is likely to be a palaeo­
forming tapering planate sprays, covered with green relict. It usually occurs, where sympatric with the
leaves; ultimate lateral branchlets partly deciduous more ubiquitous C. obtusa, on wetter sites e.g. near
after 4–6 years. Leaves decussate, imbricate, decur- mountain streams or in hollows with ground water
rent, scale-like, coriaceous, 1.5–2 × 1 mm on ultimate near the surface. In those wet places trees can spread
branchlets, up to 15 × 4 mm on leading shoots, di- by the layering of branches to form small groves of
morphic; facials smaller than laterals, rhombic to clonal individuals. It is a minor constituent in mixed
obovate, sometimes obtusely keeled, obtuse-acumi- conifer/angiosperm forests. Its altitudinal range is
nate, appressed, with a conspicous, circular, slightly from 280 m to 2600 m a.s.l.
elevated, non-active abaxial gland; laterals connate
proximally, spreading above apex of facials, condu-
Conservation
plicate, broadly falcate, recurved and incurved at
appressed apex or convex on ultimate branchlets, Although being more scattered in its (original) dis-
less conspicuously glandular; margins entire; leaves tribution than C. obtusa, this species has not been
amphistomatic, stomata concentrated on the un- exploited anywhere nearly as intensively (its timber
derside of laterals in depressed central region and value is limited) and, as a minor constituent, it sur-
on partly hidden bases and margins of leaves; leaf vives where the natural forest containing it survives.
colour lustrous light or dark green, stomatal zones IUCN: LC
glaucous white. Pollen cones numerous, terminal,
solitary, ovoid-oblong, 2–3 × 1 mm, yellowish green
Uses
turning purplish brown; microsporophylls 8–12,
decussate, peltate, subcordate, with erose margins, This species is limited as a source of timber, but
with 2–3 abaxial, yellow pollen sacs on the lower widely cultivated as an ornamental, with many
margin. Seed cones terminal on branchlets with un- named cultivars. As of other members of the genus,
modified leaves, solitary, maturing within one year, the wood is valued in Japan for traditional construc-
caducous, subglobose or irregular, 5–7(–8) mm with tion work, but its relative scarcity and the often poor
opened scales, from yellowish green (bracts) and shape of layering trees limit its commercial use.
purplish blue ripening to brown or blackish brown. However, its ornamental merits are considered sub-
Bract-scale complexes (6–)7–8(–10), decussate, stantial due to the many cultivars with different hab-
parting and spreading at varying angles from axis its and foliage (among which are forms that retain
when mature; distal pair usually connate, subpeltate juvenile type leaves) selected in Japan and Europe.
to peltate, quadrangular or irregular in outline, up Sarawa cypress is sensitive to drought and urban air
to 5 mm wide; abaxial surface centrally depressed, pollution, but is hardy in climates with not too long
rugose, sometimes with a central small umbo (bract and severe winters.
 Chamaecyparis thyoides (L.) Britton, Sterns & erose margins, with 2(–3) abaxial large, yellow pol-
Poggenb., Prelim. Cat. Anth. Pter. New York: 71. len sacs on the lower margin. Seed cones termi-
1888. nal on branchlets with unmodified leaves, solitary,
maturing within one year, caducous, subglobose,
4–7(–8) mm with opened scales, from glaucous
Etymology
purplish ­ripening to reddish brown or dark brown,
The classical Greek name for the Sandarac tree often glaucous. Bract-scale complexes (4–)6–8(–10),
(Tetraclinis articulata) or its wood was thyon, hence usually 6 in decussate pairs, parting and spreading
thyoides = similar but not equal to thyon. at varying angles from the axis when mature, pel-
tate, irregularly angular with curved margins, up
282 to 3–5 mm wide; abaxial surface depressed, rugose,
Vernacular names
with a central curved, spiny umbo (bract tip ca. 1
Atlantic White-cedar, Atlantic White Cypress, mm long); base narrowing; adaxial surface striat-
White Cypress ed, reddish brown, with faint seed marks near the
base. Seeds 2 at base of each scale (8–12 develop per
cone), slightly flattened, ovoid, shallowly grooved,
Description
with concave light coloured hilum and acutish apex,
Trees to 30–35(–40) m tall; trunk monopodial, up 2–2.5 × 2 mm, lustrous dark brown, surrounded by 2
to 1.5–2 m d.b.h. Bark soon flaky, on large trees to lateral, nearly equal wings 1 mm wide.
5 cm thick, deeply fissured and with connecting
ridges, sometimes spirally twisted from torqued
Distribution
wood, fibrous, exfoliating in long strips, reddish
brown weathering grey-brown. Branches slender, E and SE USA, from Maine south to N Florida, west
persistent, spreading or ascending; higher order to S Mississippi.
branches ascending, forming a conical crown in TDWG codes: 75 CNT MAI MAS MRY NWH NWJ
young trees, eventually becoming broader and more NWY RHO 78 ALA DEL FLA GEO MSI NCA SCA VRG
open. Foliage branches numerous, slender, drooping
or pendulous; plagiotropic branchlets alternating,
Ecology
but smallest ones more irregularly disposed, only
slightly planated, forming ‘tufts’ of foliage, covered Chamaecyparis thyoides usually grows in more
with green leaves; ultimate lateral branchlets partly or less pure stands in bogs and swamps and along
deciduous after 4–5 years. Leaves decussate, imbri- streamside corridors of lowland rivers surrounded
cate, decurrent, scale-like, coriaceous, 1.5–2.5 × 1–1.5 by other tree species, which form the main for-
mm on ultimate branchlets, up to 10 mm long on est types of the region where it occurs. Due to its
leading shoots, dimorphic, but facials on higher great latitudinal range it is associated with differ-
order branchlets only slightly shorter than the lat- ent species from N to S. The majority of these are
erals, rhombic to ovate-oblong, sometimes keeled, angiosperms, which also occupy the greater total
obtuse or acuminate, appressed, with or without a area, associated conifers are mainly Pinus spp. and
raised, active abaxial gland in the centre; laterals Taxodium distichum. The soil types under stands of
connate proximally, spreading at or less often above C. thyoides are acid organic (‘muck’) or sandy, with
the apex of facials, conduplicate, broadly falcate to the water table reaching the surface and prolonged
lanceolate, recurved and incurved at the appressed seasonal periods of inundation. It avoids salinity al-
apex, less conspicuously glandular; margins entire; though it is known to border tidal marshes in New
leaves amphistomatic, stomata inconspicuous, con- Jersey. It is likely that recurring fire would historical-
centrated on the partly hidden bases and margins of ly have been the disturbance agent preventing Acer
leaves; leaf colour yellowish green or greyish green. rubrum from eventually replacing C. thyoides in the
Pollen cones numerous, terminal, solitary, ovoid, succession. Apart from the species composition of
1.5–3 × 1–2 mm, yellowish turning light brown or the vegetation due to latitude, the ecology of the two
dark brown to blackish brown; microsporophylls varieties in this species is similar.
8–12, decussate, peltate, subcordate, with minutely
 Conservation Distribution
This species has been heavily exploited for its timber Eastern USA: coastal plain from Maine to Alabama.
and ‘total volume’ is considered to have been much TDWG codes: 75 CNT MAI MAS MRY NWH NWJ
reduced during the 20th century (Little & Garrett in NWY RHO 78 DEL FLA GEO NCA SCA VRG
Burns & Honkala, 1990). However, the species is still
very widespread (though scattered) and common in
Conservation
most swamp forests on the Atlantic Coastal Plain
and Mexican Gulf coast in Florida and Alabama, IUCN: NT
and is likely to have recovered in numbers if not yet
in ‘volume’. Chamaecyparis thyoides (L.) Britton, Sterns & 283
IUCN: For ratings see under varieties. Poggenb. var. henryae (H. L. Li) Little, Madroño 18:
165. 1966. Chamaecyparis henryae H. L. Li, Morris
Arbor. Bull. 13 (3): 43. 1962; Chamaecyparis thyoides
Uses
(L.) Britton, Sterns & Poggenb. subsp. henryae
The wood of this species is light, decay resistant, and (H. L. Li) E. Murray, Kalmia 12: 19. 1982. Type: USA:
is still widely used in the SE USA for many purposes Florida, Escambia Co., Perdido River, at Barineau
involving outdoor utilities. Trees are occasionally Park, M. G. Henry 23 (holotype PH).
cultivated and sold as ornamentals but there is no
substantial horticultural trade. A limited number
Description
of cultivars is known and most of the plants in cul-
tivation belong to one of these. Among these are Scale leaves on ultimate branchlets eglandular or
some dwarf forms and one of these, ‘Andelyensis’ inconspicuously glandular, facials often distinct-
has incorrectly been described as a botanical variety ly keeled abaxially. Pollen cones at maturity light
by the German horticultural botanist Camillo Karl brown.
Schneider. Forms originating in cultivation are not
botanical varieties (or species) and under the rules
Distribution
of horticultural nomenclature are now to be given
non-latinized cultivar names. SE USA: along the Gulf Coast from Florida to Mis-
sissippi.
2 varieties are recognized: TDWG codes: 78 ALA FLA MSI

Chamaecyparis thyoides (L.) Britton, Sterns &

Conservation
Poggenb. var. thyoides. Cupressus thyoides L., Sp.
Pl. 2: 1003. Type: USA: [locality unknown], P. Kalm IUCN: NT
LINN 1137.4 (lectotype LINN). Fig. 90

Description
Scale leaves on ultimate branchlets conspicuously
glandular, especially facials, these not keeled. Pollen
cones at maturity dark brown or blackish brown.
 Cryptomeria D. Don, Ann. Nat. Hist. 1: 233. 1838. Type: Cryptomeria japonica
(Thunb. ex L. f.) D. Don [Cupressus japonica Thunb. ex L. f.] (Cupressaceae).

Greek: kryptos = covered, hidden; -meros = share strips. Branches spreading to assurgent, forming a
(parts or their number); referring to seeds hidden conical crown in young trees and a rounded crown
by bracts. in mature trees, self-pruning to leave a clear bole in
large trees. Foliage branches dense, shedding not in-
dividual leaves but ultimate lateral branches which
Description
persist 4–8 years. Leaves helically arranged in ranks
284 See the species description. of 5, decurrent, free for 1/2–3/4 of length, spreading
but incurved in various degrees, directed forward,
linear-subulate, slightly flattened laterally, distinctly
Distribution
keeled abaxially, stiff, green, 3–20(–25) × 1–2 mm;
As for the species. margins entire; apex acute; amphistomatic, stomata
in 4 greenish bands separated by green ribs, usually
on entire length of leaf. Pollen cones numerous, axil-
Cryptomeria japonica (Thunb. ex L. f.) D. Don, lary and crowded towards ends of 2nd-year branch-
Trans. Linn. Soc. London 18: 167. 1839. Cupressus lets, 3–6 × 2–3 mm, elongating to 10 mm at anthesis;
japonica Thunb. ex L. f., Suppl. Pl.: 421. 1781. microsporophylls 15–30, the first 2 opposite and
Type: Japan: Honshu, [“Habitat in Japonia”], green, then helically arranged, imbricate, peltate,
C. P. Thunberg UPS 22564 (lectotype UPS). Fig. 91 with 4–6 connate pollen sacs on the lower abax-
ial margin. Seed cones terminal on down-curved
Cryptomeria fortunei Hooibr., Wiener J. Gesammte branchlets with normal leaves, often aggregated or
Pflanzenr. 1: 22. 1853, nom. nud. [“fortunini”]; Cryp- solitary, occasionally with proliferating vegetative
tomeria japonica (Thunb. ex L. f.) D. Don var. for- short shoot at apex, globose to subglobose, squar-
tunei (Hooibr.) Henry, in Elwes & Henry, Trees Gr. rose with spreading bract-scale complexes, soft
Brit. Ireland 1: 129. 1906 (nom. inval., Art. 34.1). woody, 12–20(–25) mm diam. Bract-scale complexes
Cryptomeria japonica (Thunb. ex L. f.) D. Don var. helically arranged on a very short axis, 25–40, con-
sinensis Miq., in Siebold & Zuccarini, Fl. Japon. 2: 52. nate but parting at maturity, cuneate-rhombic, nar-
1870; Cryptomeria japonica (Thunb. ex L. f.) D. Don rowly based; apex a curved, triangular, thin bract;
subsp. sinensis (Miq.) P. D. Sell, Watsonia 18 (1): 92. teeth adaxial to the bract below the upper margin of
1990. the scale, (1–)3–5(–6) in number, acute to rostrate,
(2–)2.5–5(–6) mm long. Seeds (1–)2–5 per bract-
scale complex (some ovules may abort) depending
Etymology
on space available when intercalary scale tissue de-
The species epithet refers to Japan, the country of velops, 4–5 × 3 mm, flattened, irregularly ovate with
origin of this species. 2 wings; wings unequal, 1–1.5 mm wide, forming a
strip around the seed.
Vernacular names
Taxonomic notes
Japanese cedar; Sugi, Omote-sugi, Yaku-sugi, (Japa-
nese) Well before Cryptomeria was introduced into Eu-
rope in 1842 (England), there were two provenances
known: China and Japan. European knowledge of
Description
the Japanese trees goes back to Engelbert Kaempfer
Trees to 50–60 m tall, evergreen, monoecious; trunk (1690–92), that of the Chinese trees to James Cun-
straight, columnar, usually monopodial, in old trees ningham (1701–02). They have long been regarded
massive, buttressed, 4–5 m (formerly to 8 m) diam. as two distinct species, or sometimes as different
Bark on large trees 2–3 cm thick, reddish brown, varieties, but the minor morphological differ­ences
weathering grey, exfoliating in long, shredding ascribed to the Chinese taxon can be found in
s­ pecimens that originate from Japan as well. The ­ ydrangea petiolaris and Rhus orientalis; a rich cryp-
H
opposite is also the case and in conclusion all one togamic flora covers the forest floor as well as trees,
can say is that plants from China more often have with abundant ferns, e.g. Hymenophyllum, mosses,
short ‘teeth’ and plants from Japan usually have and liverworts. The mountains on this island are of
longer ‘teeth’ on the seed cones. The explanation granite; the soils are well drained yellow loam or
for this is probably that the short-toothed form in clay, often quite deep. The climate is mild temperate,
China is based on an early introduction of limited with abundant rainfall.
provenance with that cone form; later repeated in-
troductions came from a wider range of trees and
Conservation
included the more common form with longer teeth.
There can be little doubt that Cryptomeria was in- Extensive logging has removed trees of this species 285
troduced long ago to China (Farjon, 1999), and that in much of its natural range and few old growth for-
its inclusion in Floras and other accounts is based ests with Cryptomeria remain. On the other hand,
on uncritical assessment of data on distribution de- plantation forestry in Japan has made extensive use
spite earlier doubts expressed by Wilson (1926) and of the species, bringing it back to many areas where
Florin (1963). Even in Japan, where Cryptomeria is it had been greatly depleted. The species is undoubt-
endemic, the natural distribution is apparently now edly regenerating naturally from this stock in many
very difficult to establish. This species has not been forests, so that the distinction between its natural
mapped in the Atlas of the Japanese Flora (Hori- distribution and anthropogenic occurrences can
kawa, 1972), presumably because of these problems. only be ascertained by a detailed study of forest his-
tory in Japan that would distinguish between pri-
mary and secondary occurrence.
Distribution
IUCN: NT
Japan: Hokkaido (?), Honshu, Kyushu, Shikoku,
Yakushima.
Uses
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
The timber of this species is regarded in Japan as be-
ing highly valuable for construction because of its
Ecology
size, workability, and durability. This species is the
Natural forests that include this species are now very principal conifer in Japanese forestry and almost
rare (Tomaru et al., 1994) and those forests in which all timber now comes from plantations. It has been
it still occurs have been greatly altered; the descrip- introduced to China many centuries ago; in Tai-
tion is largely based on observations by E. H. Wilson wan the Japanese established extensive plantations
made in 1914 on Yakushima (Wilson, 1916), where replacing natural forest. The wood is rather coarse
an old growth forest still exists. The forest vegetation grained, with reddish heartwood and light yellow-
is mixed evergreen forest, with ca. 50% Crypto­meria, ish sapwood, and a typical coniferous fragrance due
growing mixed or in groves; angiosperm evergreen to volatile terpines. It is used in house construction,
trees are Trochodendron aralioides, ­Distylium race- panelling, flooring, carpentry, joinery, and furniture
mosum, Camellia japonica, C. sasquana, Daphni­ making. The fibrous bark was traditionally used for
phyllum spp., Michelia compressa, Myrica rubra, roofs of houses but is now more often processed as
Quercus spp., Ilex spp., and Lauraceae; conifers are a general fibre or composted. There are several hun-
Abies firma, Tsuga sieboldii, Chamaecyparis obtusa, dred cultivars known of this species, in Japan and in
and Torreya nucifera; a few deciduous angiosperm Europe and North America, and the species as well
trees, e.g. Stewartia monadelpha and Acer spp., as many of these cultivars are widely used in gardens
make up less than 1% of tree cover. There is a di- and parks.
verse shrub-layer and some common climbers, e.g.
 Cunninghamia R. Br., in Richard, Conif. Cycad.: 80, 149. 1826 (nom. cons.). Type:
Cunninghamia lanceolata (Lamb.) Hook. [Cunninghamia sinensis R. Br. (nom. illeg.)
(Pinus lanceolata Lamb.)] (Cupressaceae).

Named after Allan Cunningham (1791–1839), Eng-

Key to the species of Cunninghamia
lish traveller and botanist.
1a. Leaves 12–30 × 2–3 mm, conspicuously amphis-
tomatic; adaxial stomata in continuous narrow
Description
bands from base to apex of the leaf C. konishii
286 Evergreen, monoecious trees; trunk monopodial, 1b. Leaves 30–60 × 3–5(–6) mm, hypostomatic, or
readily resprouting. Resin ducts in leaves. Bark fis- at most with a few intermittent or incomplete
sured, fibrous, exfoliating in long strips, reddish lines of stomata adaxially C. lanceolata
brown. Branches in whorls, long, slender, higher or-
der branches spreading or drooping to pendulous,
plagiotropic (Massart’s model); capacity to cop- Cunninghamia konishii Hayata, Gard. Chron., ser.
pice profound. Foliage branches slender, branch- 3, 43: 194. 1908. [J. Linn. Soc., Bot. 38: 299. 1908].
ing (nearly) opposite, lateral branchlets deciduous. Type: Taiwan: Chiayi Co., Luan-ta Shan, [“Mt.
Leaves helically arranged, decurrent at distinctly Rantaizan”], N. Konishi s.n. (holotype TI). Fig. 92
narrowed base, free part curved or twisted into a
more or less pectinate arrangement with primary
Etymology
stomatal face on underside, coriaceous, flattened,
linear-­lanceolate, gradually tapering to an acute The species epithet commemorates N. Konishi, the
apex; margins serrulate; amphistomatic or hypo­ collector of the type specimen.
stomatic; abaxial stomata in 2 broad glaucous white
bands separated by a green elevated midrib; adax-
Vernacular names
ial stomata in two narrow bands (3–5 lines) along
margins or absent. Pollen cones subterminal on fo- Taiwan shan mu (Chinese)
liage branches, numerous in clusters subtended by
a pseudo-whorl of bract-like short leaves, cylindri-
Description
cal; microsporophylls numerous, helically arranged,
peltate, bearing 3(–4) abaxial pollen sacs. Seed cones Trees to 40–50 m tall; trunk monopodial, often but-
subterminal, solitary or paired, persistent (falling tressed in old trees, readily resprouting (coppice
with foliage branches); mature cones ovoid-globose. or pollard trees), to 3(–4.5) m d.b.h. Bark on trunk
Bract-scale complexes numerous, helically arranged, fissured, fibrous, exfoliating in long strips, reddish
imbricate, appressed at base, spreading distally at brown weathering dull brown. Branches in whorls,
maturity, more or less triangular with a pedicel- long, slender, spreading to ascending near the top,
late base and an apiculate to cuspidate apex, coria- forming a pyramidal to finally irregular, rounded
ceous; abaxial surface smooth; adaxial surface with crown; higher order branches spreading or drooping
2–3 seed marks at the distal end of the seed-bearing to pendulous, 3rd and 4th order branches plagiotropic
tissue. Seeds 2–3 per fertile scale, with 2 marginal, (but profuse reiteration after damage obscures this
1 mm wide wings. Seedlings with 2 cotyledons. arrangement). Foliage branches numerous, slen-
der, branching (nearly) opposite, lateral branchlets
2 species. deciduous. Leaves decurrent at distinctly narrowed
base, free part curved or twisted into a more or less
pectinate arrangement with the primary stomatal
Distribution
face on the undersides, flattened, narrowly lanceo-
China, Laos, Taiwan, Vietnam. late or linear-lanceolate, straight or s-curved, gradu-
ally tapering to an acute apex; margins serrulate,
12–30 × 2–3 mm; amphistomatic, abaxial (lower)
Distribution
stomata in 2 broad glaucous white bands of 10–20
irregular lines separated by a green elevated mid- Taiwan (N-central); China, Fujian; Laos, Houaphan
rib; adaxial stomata in two narrow bands (3–5 lines) Province; Vietnam (Bu Huong Mountain, Phu Hoat
along margins; adaxial surface smooth, glaucous Mountain).
green or green. Pollen cones subterminal on foliage TDWG codes: 36 CHS-FJ 38 TAI 41 LAO VIE
branches, in clusters subtended by a pseudo-whorl
of bract-like short leaves, 10–15 × 3–4 mm; micro-
Ecology
sporophylls 50 or more, helically arranged on a slen-
der axis, peltate; margins erose-denticulate; apex In mixed coniferous or conifer-broad-leaved for-
acute; bearing 3 abaxial, large, oblong pollen sacs on est in the cool temperate coniferous forest belt, 287
the lower margin. Seed cones subterminal, solitary with Chamaecyparis formosensis, C. obtusa var.
or paired, maturing within 1 year, persistent (fall- formosana, Calocedrus formosana, Pinus taiwanen-
ing with foliage branchlets); mature cones ovoid- sis, Pseudo­tsuga sinensis, Taiwania crypto­merioides,
globose, 15–25 × 15–20 mm, turning lustrous brown. Acer morrisonense, A. kawakamii, Schima ­superba,
Bract-scale complexes numerous, helically arranged Photinia davidiana, Rhododendron ­formosanum,
on a thin axis, persistent, imbricate, appressed at Pasania sp., and the alpine bamboo Yushania nii-
base, spreading distally at maturity, more or less tri- takayamensis. The altitudinal range is from (600?)
angular with a pedicellate base, coriaceous; margins 1000 m to 2200 m a.s.l. Soils are relatively deep and
entire or denticulate near apiculate apex; abaxial well-drained loams or loamy sand. The climate is
surface smooth, more or less keeled towards apex. cool, very moist, with cloud-cover resulting in fog
Seeds usually 2 per fertile scale, obovate, flat, 4–6 × and rain much of the year and very high precipita-
3–4 mm, brown with a light hilum near the base and tion exceeding 4000 mm per year.
2 marginal wings 1 mm wide leaving an emarginate
seed apex. Teratism in the form of ‘shoot prolifera-
Conservation
tion common.
This species is now known from very disjunct locali-
ties in warm temperate E Asia, but outside Taiwan
Taxonomic notes
its conservation status, and indeed its areas of occu-
Lu et al. (1999) investigated molecular data from a pancy and abundance, are poorly known. In Taiwan,
small part of the chloroplast genome (trnD-trnT substantial decline has occurred in the populations
spacer) in four samples from Taiwan and four from due to felling of original forests, which are being re-
mainland China and found the sampled populations placed by plantations if not converted to other land
to be paraphyletic. This of course means in evolu- uses. In those cases where Cunninghamia has been
tionary terms that this still extant species may have replanted, use has mostly been made of C. lanceo-
been the ancestor of its also extant congener. It does lata, which shows faster growth than. C. konishii.
not mean that C. konishii and C. lanceolata can- This poses another threat: genetic contamination.
not be distinct species, unless one insists that spe- A magnificent stand of giant trees of C. konishii is
cies be monophyletic, which this author does not. protected in Taroko National Park, far from any
Phylogenetic inference from such a limited sample planted trees.
base is questionable anyway. In addition, there is a IUCN: VU (A1c)
compounding problem with the provenance of es-
pecially C. lanceolata, as few trees in China are truly
Uses
originating from wild populations in undisturbed
forests. In contrast, C. konishii is known from re- The timber of this species is valued for its durabil-
mote forest sites, at least in Taiwan, Laos, and Viet- ity under wet climate conditions. As a rare tree its
nam. If more becomes known about these and other, commercial exploitation is now negligeable, but in
disjunct populations of C. konishii, it is possible that the past is was logged and large trees were very valu-
we must conclude that it is a montane form of a able. In cultivation it is uncommon, mainly restrict-
widespread, polymorphic species, or perhaps that ed to botanical collections in arboreta and botanic
we are dealing with a few ‘cryptic’ species. ­gardens.
 Cunninghamia lanceolata (Lamb.) Hook., Bot. to acute; bearing 3(–4) abaxial, large, oblong pol-
Mag. 54: sub t. 2743. 1827. Pinus lanceolata Lamb., len sacs on lower margin. Seed cones subterminal,
Descr. Pinus 1: [52], t. 34. 1803. Type: Illustration solitary or in clusters of 2-several, maturing within
in Lambert, Descr. Pinus 1, t. 34. 1803 (lectotype). 1 year, persistent (falling with foliage branches); ma-
Fig. 93 ture cones ovoid-globose, 25–40 × 25–35 mm, turn-
ing lustrous reddish brown. Bract-scale complexes
Cunninghamia unicanaliculata D. Y. Wang & spirally arranged on a thin axis, persistent, imbri-
H. L. Liu, Acta Phytotax. Sin. 20 (2): 230. 1982. cate, appressed at base, spreading distally at matu-
Cunninghamia unicanaliculata D. Y. Wang & rity, more or less triangular with a pedicellate base,
H. L. Liu var. pyramidalis D. Y. Wang & H. L. Liu, coriaceous; margins denticulate; apex cuspidate to
288 Acta Phytotax. Sin. 20 (2): 231. 1982. rostrate; abaxial surface smooth, more or less keeled
towards apex; adaxial surface with 2–3 seed marks at
the distal end of the seed-bearing tissue. Seeds usu-
Etymology
ally 2 per fertile scale, obovate, flat, 6–7 × 4–5 mm,
The species epithet refers to the shape of the leaves brown with a light hilum near base and 2 margin-
resembling the point of a lance. al wings 1–2 mm wide leaving an emarginate seed
apex. Teratism in the form of ‘shoot proliferation
frequent.
Vernacular names
Chinese fir; shan mu (Chinese)
Taxonomic notes
The new species C. unicanaliculata and its variety
Description
pyramidalis (the latter based on a tree with slender
Trees to 30–40 m tall; trunk monopodial, often but- habit), described by Wang & Liu (op. cit.) were sunk
tressed in old trees, readily resprouting (coppice or into synonymy in Flora of China 4 (1999). Wang &
pollard trees), up to 2.5–3(–6?) m d.b.h. Bark on Liu stated that the new species differs from C. kon-
trunk fissured, fibrous, exfoliating in long strips, ishii by “frequent” single resin canals in the leaves,
reddish brown weathering to dull brown. Branches but there is no indication of what higher numbers, if
in whorls, long, slender, spreading to ascending near any, they observed in other leaves belonging to Cun-
the top, ), forming a pyramidal to finally irregular, ninghamia. Both C. konishii and C. lanceolata nor-
rounded crown; higher order branches spreading mally have a single resin canal in the leaves.
or drooping, 3rd and 4th order branches plagiotropic
(but profuse reiteration after damage obscures this
Distribution
arrangement). Foliage branches numerous, slender,
branching (nearly) opposite; lateral branchlets de- China: from Sichuan to the coast, mainly in SE Chi-
ciduous. Leaves decurrent at slightly narrowed base, na; Laos, Vietnam (doubtfully indigenous).
free part curved or twisted into a more or less pecti- TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
nate arrangement with stomatal face on underside, SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS-
flattened, lanceolate-linear, straight or curved, grad- HE CHS-HN CHS-JS CHS-JX CHS-ZJ 41 LAO VIE
ually tapering to a pungent apex; margins serrulate;
30–60 × 3–5(–6) mm; hypostomatic, with stomata
Ecology
in 2 broad glaucous white bands of 10–35 irregular
lines separated by a green slightly elevated mid- Wang (1961) considered C. lanceolata to be a coni­
rib; adaxial surface smooth, with 2 shallow lateral ferous constituent of the Mixed Mesophytic Forest
grooves, lustrous green. Pollen cones (sub)terminal Formation of the warm temperate regions of Chi-
on foliage branches, numerous in clusters subtended na. This is predomiantly a deciduous angiosperm-­
by a pseudo-whorl of bract-like short leaves, 10–20 × dominated forest in which no particular species
3–5 mm, yellowish green turning brown; microspo- dominate and conifers form a minor but fairly con-
rophylls 50 or more, spirally arranged on a slender stant component. Almost all of the original forest
axis, peltate; margins erose-denticulate; apex obtuse has disappeared under the long-lasting influence of
 a dense rural population. In remnants of natural veg-
Conservation
etation, e.g. Tienmu Shan in Zhejiang, it occurs with
numerous species in Acer, Magnolia, Quercus, and IUCN: LC
other angiosperms. The conifers mentioned for this
mountain are in part natural understorey species of
Uses
this forest type, i.e. Cephalotaxus fortunei, Pseudo-
taxus chienii, Taxus chinensis, and Torreya grandis. Uses of this tree vary, from timber to firewood and
However, most of the other conifers mentioned to horticultural, with amenity planting in Chinese cit-
occur in it are either introduced in Zhejiang (Crypto­ ies and parks as well as botanical collections and
meria japonica (Farjon, 1999), ­Cupressus funebris, gardens in other countries. The wood is light, soft,
or not typical for old growth of this forest type, but fragrant, almost white, and durable. Large sizes are 289
for secondary vegetation (e.g. Juniperus formosana, milled for construction timber in houses, for masts,
­Pinus massoniana, and P. ­tabuliformis). Many habitat carpentry and planks for coffins. In the past, wood
notes on herbarium sheets of C. lanceolata mention buried in landslides was reported to be dug up and
forest, forest edges, light woodland, rocky slopes, as it was darker, was much valued for coffins. In Eu-
and scrub, most of which indicate secondary vegeta- rope a few cultivars, primarily producing dwarfed
tion. The capacity to coppice and relative intolerance habits, have been selected but the tree is most com-
to shade will ensure its survival in cutover forest ar- monly seen in arboreta as the species. In China ad-
eas. It is also widely planted in afforestation projects vantage is taken of its capacity to coppice, i.e. grow
throughout warm temperate China and beyond. new stems from cut tree stumps and roots, and
much of this is used as firewood.
 Cupressus L., Sp. Pl. 2: 1002. 1753. Type: Cupressus sempervirens L. (Cupressaceae).

Tassilicyparis A. V. Bobrov & Melikyan, Komaro- 15 species.

via 4: 72. 2006. Type: Tassilicyparis dupreziana
(A. Camus) A. V. Bobrov & Melikyan [Cupressus
Distribution
dupreziana A. Camus]. Platycyparis A. V. Bobrov &
Melikyan, Komarovia 4: 73. 2006. Type: Platycyparis SW North America, S to Honduras; North Africa
funebris (Endl.) A. V. Bobrov & Melikyan [Cupressus and Mediterranean to Middle East; Himalaya to SW
funebris Endl.]. Hesperocyparis Bartel & R. A. Price, China.
290 Phytologia 91 (1): 179. 2009. Type: Hesperocyparis
macrocarpa (Hartw. ex Gordon) Bartel [Cupressus
Taxonomic notes
macrocarpa Hartw. ex Gordon].
In a recent paper, Little (2006) used (mainly) DNA
Cupressus is the classical Latin name for cypresses. nucleotide sequence data in a phylogenetic analy-
sis and followed the cladistic paradigm to classify
the species. Juniperus appeared to be derived from
Description
(‘nested within’) Cupressus. To avoid paraphyly of
Shrubs or trees to 40(–95) m, evergreen, monoe- Cupressus, the genus had therefore to be split into
cious; trunk usually monopodial, sometimes but- two (the alternative under cladisitic insistence on
tressed. Resin cavities in leaves. Bark smooth or monophyletic taxa would be to sink Juniperus with
more commonly fissured and fibrous, hard or soft, its more than 50 species into Cupressus). In a re-
exfoliating in small flakes or long strips. Branches sponse it was argued (Farjon, 2007) that it is pos-
long and spreading or ascending, eventually droop- sible to recognize that Juniperus might indeed have
ing or pendulous in several species, forming coni- evolved from an ancestor which, had we known it,
cal, pyramidal or rounded crowns. Fastigiate forms would be classified as a species of Cupressus based
mostly restricted to cultivation. Foliage branches ir- on the morphological circumscription of that genus.
regularly disposed and spreading or plagiotropic and Evolutionary (phyletic) classification accepts but
pendulous, also intermediate forms. Leaves scale- does not insist on Hennigian monophyletic (= holo­
like, monomorphic or dimorphic, appressed or with phyletic) taxa. Most recently, another cladistic
free apices, on ultimate branchlets 1–3 × 0.8–1.2 mm, analysis using molecular data (Adams et al., 2009)
on leading shoots larger, triangular to rhombic or, gave similar results, but with the New World species
in branchlets with dimorphic leaves, with laterals of Cupressus now (again) separated from the two
slightly larger than facials and conduplicate, glan- species of Xanthocyparis as here recognized (these
dular or eglandular; margins minutely denticulate were included in the New World Cypresses in Lit-
or erose-hyaline, sometimes entire. Pollen cones tle’s 2006 paper, but not in an earlier analysis based
terminal, solitary, 3–7 × 2–3 mm, with 10–20 dec- on nuclear ITS data). Little had ‘resurrected’ the
ussate microsporophylls bearing 3–6 abaxial yellow forgotten name Callitropsis Oerst. for all these and
pollen sacs. Seed cones solitary on short branchlets, made the appropriate new combinations, but that
often grouped close together or clustered, cadu- name has subsequently been proposed and accepted
cous or persistent, globose to ovoid-oblong 10–40 (subject to ratification at the International Botani-
× 10–30 mm, with parting woody scales. Bract- cal Congress in 2011) for official rejection because of
scale complexes in (3–)4–7 decussate pairs, peltate, ambiguity and other reasons causing disruption of
quadrangular, rhombic or polygonal in outline, with names. So, again adhering to the cladistic principle
well developed umbos (bract apices) in most cases; of monophyly, a new name was now proposed for
adaxially with light seed marks towards narrowing the New World Cypresses (sensu stricto): Hespero­
base. Seeds numerous, in two or more rows on bases cyparis Bartel & R. A. Price, with the appropriate
of scales, small, angular, with rudimentary wings. new combinations. It is admitted by these authors,
Cotyledons 2–5, juvenile leaves only on seedlings, in that “the new genus is cryptic in its macromorphol-
whorls of 3–4, acicular-linear. ogy”, although unconvincing attempts were made to
list some differences. Very few of these differences
apply to Cupressus macrocarpa (the type of the new 7b. Seeds with equal wings, brown or blackish
genus Hesperocyparis) and C. sempervirens (the type brown, not glaucous C. macrocarpa
of Cupressus) and if taken across the genus Cupres- 8a. Pollen sacs 2–3 per microsporophyll. American
sus as commonly circumscribed, none of them are species 9
consistent. Another contentious issue appears to be 8b. Pollen sacs 4–5, if 2–3, then Eurasian / North
the generic placement of the Leyland Cypress (as African species 11
such not treated in this Handbook as it is not a taxon 9a. Bark smooth, exfoliating in numerous, papery
that occurs in the wild). If its parents both belong to flakes C. bakeri
Cupressus, then it is not an intergeneric hybrid, but 9b. Bark becoming fissured, hard fibrous 10
just a hybrid between two species of that genus. Both 10a. Seed cones 15–30 mm diam, often irregular in
parents, Monterey cypress and Nootka cypress, have shape. Californian species C. sargentii 291
been removed from Cupressus (the first by Little and 10b. Seed cones 10–18(–20) mm diam., usually glo-
by Adams et al., cited above, the latter in the 19th cen- bose and regular in shape. Species distributed
tury to Chamaecyparis and recently to Callitropsis from Mexico to Honduras C. lusitanica
and, as accepted here, to ­Xanthocyparis). The con- 11a. Foliage branchlets slender, lax 12
troversy here is not purely one of taxonomy; it has 11b. Foliage branchlets thick, rigid 14
a direct bearing upon the (correct) botanical name 12a. Seeds fewer than 10 on each cone scale 13
of this economically important cultivated conifer. 12b. Seeds more than 10, at least on the middle cone
Horticulturists could be tempted, after reading all scales C. duclouxiana
of this, to forgo on botanical names altogether; to 13a. Seed cone scales with prominent, curved boss-
them the Leyland Cypress will remain the Leyland es caused by protruding bract tips C. torulosa
Cypress. 13b. Seed cone scales with small, inconspicuously
protruding bract tips C. chengiana
14a. Seed cones 15–27 × 13–21 mm. North African
Key to the species of Cupressus
species C. dupreziana
1a. Leaves on ultimate branchlets dimorphic; later- 14b. Seed cones 20–35 × 20–25 mm. Species distrib-
als larger than facials; glands present on facials uted in E Mediterranean and Middle East
2 C. sempervirens
1b. Leaves on ultimate branchlets all of similar
shape and size, or if dimorphic, eglandular 3
2a. Seed cones with 6–8 scales. Lateral leaves usu- Cupressus arizonica Greene, Bull. Torrey Bot. Club
ally with appressed apex C. funebris 9: 64. 1882.
2b. Seed cones with (8–)10 scales. Lateral leaves
with free apex C. cashmeriana
Etymology
3a. Seed cones usually in dense, serotinous clusters.
Pollen sacs often 5 or more per microsporo­ The species epithet refers to the US State of Arizona
phyll 4 where this species is native.
3b. Seed cones solitary or in groups, not densely
clustered. Pollen sacs up to 4 per microsporo-
Vernacular names
phyll 8
4a. Glands present on scale leaves 5 Arizona cypress; Cedro, Cedro blanco (Spanish)
4b. Leaves, at least those of ultimate branchlets, eg-
landular 6
Description
5a. Seed cones with 6, rarely up to 8 scales
C. macnabiana Large shrubs or trees 5–20(–30) m tall; trunk mo-
5b. Seed cones with 8 or more scales C. arizonica nopodial or multistemmed from near the ground,
6a. Bark remaining smooth and scaly even on very to 40–100 cm d.b.h. Bark smooth on branches and
large shrubs or trees C. guadalupensis younger trees, soon turning grey, then exfoliating
6b. Bark becoming fissured, hard fibrous 7 with thin scales and strips, exposing cherry-red
7a. Seeds with rudimentary wings or unequal or purplish red bark, remaining smooth on large
wings, brown, often glaucous C. goveniana stems (except eventually near base) and shaggy with
 e­ xfoliating thin strips, or becoming rough, scaly or Monograph of Cupressaceae and Sciadopitys (Far-
firbrous, to 4 cm thick, fissured, breaking into ir- jon, 2005a).
regular, small flakes or long, fibrous strips, brown
or blackish brown. Branches long and spreading, or
Distribution
more often ascending or erect, forming a conical,
broad conical, pyramidal, or oval and round-topped SW USA: California, Arizona, New Mexico, Texas;
crown. Foliage branches spreading or ascend- Mexico: Baja California Norte, Sonora, Chihuahua,
ing, forming tufts of coarse, rigid foliage, ultimate Coahuila, Durango, Zacatecas, Nuevo León, San
branchlets numerous, spreading outward at 45–80 Luis Potosí, Tamaulipas.
degrees, (5–)8–12(–20) mm long, quadrangular in TDWG codes: 76 ARI CAL 77 NWM TEX 79 MXE-
292 cross-section, 1.3–1.6(–2) mm diam., persistent. CO MXE-CU MXE-DU MXE-NL MXE-SL MXE-TA
Leaves decussate, decurrent, imbricate, appressed MXE-ZA MXN-SO
or on older branchlets and whip shoots several with
spreading apex, of equal size and shape on ultimate
Ecology
branchlets, triangular-rhombic, more or less gibbous
or more often with a central depression, (1–)1.4–2 In montane coniferous forest, mixed broad-leaf-
× 1–1.3 mm (much larger on whip shoots); mar- coniferous woodland, Pinyon-Juniper woodland,
gins hyaline-serrulate or denticulate; apex obtuse sclerophyllous scrubland (‘chaparral’), and valley
to acute; stomata on the abaxial side near base, on scrub-grassland. The altitudinal range is between
the adaxial side scattered from base to apex; glands 750 m and 2700 m a.s.l. Associated species in these
central, circular, conspicuous and active with a resin vegetation types are Pseudotsuga menziesii, Abies
drop, or inconspicuous and inactive, or absent on concolor, Calocedrus decurrens, Pinus ponderosa,
some branchlets; leaf colour variable, dull green, P. jeffreyi, P. arizonica, P. leiophylla, P. lambertiana,
grey-green, glaucous green, in young trees often P. sabiniana, P. coulteri, P. cembroides, P. edulis,
strongly glaucous, often covered with thick cuticular P. monophylla, Juniperus deppeana var. pachyphlaea,
wax. Pollen cones numerous, terminal, solitary, sub- J. californica, Fraxinus velutina, Quercus spp., Gar-
globose, elliptic-oblong or cylindrical, 3–5 × 1.7–2.2 rya sp., Cercocarpus sp., Platanus sp., Populus tremu-
mm; microsporophylls (8–)12–14(–16), decussate, loides, Salix sp. (along creeks), Rhus ovata, Fremontia
peltate, each bearing abaxially (2–)3–5(–6) subglo- californica, Yucca whippleyi, Adenostoma fascicula-
bose-angular pollen sacs. Seed cones terminal, of- tum, Arctostaphylos glandulosa, Ceanothus spp., and
ten grouped or clustered, maturing in two growing Rhamnus sp. It is usually ‘gregarious’ and occurs on
seasons, serotinous or opening within a year, sub- ridges, slopes and in canyons, sometimes in creek
globose to irregularly broad ovoid, 15–27(–30) mm, beds, in rocky terrain in yellow or red-brown loam,
while growing with prominent bosses on the scales. sand or gravel, or among boulders over limestone,
Bract-scale complexes decussate, (6–)8–10(–12), sandstone, slate or granite. The climate is character-
with 1–2 basal pairs usually fused or connate, the ized by warm to hot, dry summers and winter rain-
others parting at maturity, peltate, the distal part fall.
irregularly 4–6 sided, smooth, becoming coarsely
rugose, with a subapical to central, curved boss 2–6
Uses
mm long formed by the partly emerging bract, prox-
imal part abruptly constricted, pedicellate, grooved, There are no known commercial uses of the wood at
with light seed scars towards base. Seeds 90–120 per the present time. The species, in particular var. gla-
cone, irregularly orbicular or oval, (3–)4–5(–6) mm bra from Arizona, has for a considerable time been
long, more or less flattened; hilum (nearly) basal; cultivated in gardens and parks in Europe and the
wings 2, ca. 1 mm wide, surrounding the seed. USA as an ornamental. Young trees have a naturally
conical habit and several cultivars have been select-
ed to enhance this shape; other cultivars emphasize
Taxonomic notes
juvenile leaf type and extreme glaucousness of fo-
A discussion of the taxonomy of the Cupressus ari- liage. This species is tolerant of draught and grows
zonica ‘complex’ with comprehensive reference to well on chalk in full sun.
the literature on this subject can be found in the
5 varieties are recognized:
Distribution
Cupressus arizonica Greene var. arizonica. Calli­ USA: Arizona (Coconino, Gila, Maricopa and Yava-
tropsis arizonica (Greene) D. P. Little, Syst. Bot. 31 pai Co.).
(3): 473. 2006 (nom. ut. rej., Art. 56); Hesperocyparis TDWG codes: 76 ARI
arizonica (Greene) Bartel, Phytologia 91 (1): 180.
2009. Type: USA: Arizona, Greenlee Co., Clifton,
Conservation
[“on the mountains back of Clifton”], E. L. Greene
s.n. (holotype NA). Fig. 94 IUCN: LC

293
Description Uses
Bark becoming fissured or fibrous on larger trunks, This variety is in cultivation as a decorative tree for
to 4 cm thick, exfoliating slowly in small chips or large gardens and parks and is valued for its decora-
shreddy strips. Leaves inconspicuously glandular tive smooth bark and attractive green or glaucous
or eglandular, or occasionaly conspicuously and ac- foliage.
tively glandular. Boss on cone scales usually large.
Seeds red-brown to dark brown, not glaucous. Cupressus arizonica Greene var. montana (Wig-
gins) Little, Madroño 18: 163. 1966. Cupressus mon-
tana Wiggins, Contr. Dudley Herb. 1 (5): 161. 1933;
Distribution
Calli­tropsis montana (Wiggins) D. P. Little, Syst.
USA: Arizona, New Mexico, W Texas (Chisos Mts.); Bot. 31 (3): 474. 2006 (nom. ut. rej., Art. 56); Hespe-
N and NE Mexico: Chihuahua, Coahuila, Durango, rocyparis montana (Wiggins) Bartel, Phytologia 91
Sonora, Tamaulipas, Zacatecas. (1): 182. 2009. Type: Mexico: Baja California Norte,
TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CU Sierra San Pedro Martír, La Encantada, I. L. Wiggins
MXE-CO MXE-DU MXE-TA MXE-ZA MXN-SO &; D. Demaree 4990 (holotype DS).

Conservation Description
IUCN: LC Bark on trunks becoming fissured, exfoliating slow-
ly in shreddy flakes and strips. Leaves conspicuously
Cupressus arizonica Greene var. glabra (Sudw.) and actively glandular. Boss on cone scales usually
Little, Madroño 18: 162. 1966. Cupressus glabra small and obtuse. Seeds red-brown or dark brown,
Sudw., Amer. Forestry 16: 88. 1910; Callitropsis not glaucous.
glabra (Sudw.) D. P. Little, Syst. Bot. 31 (3): 473.
2006 (nom. ut. rej., Art. 56); Hesperocyparis
Distribution
glabra (Sudw.) Bartel, Phytologia 91 (1): 181. 2009.
Type: USA: Arizona, Yavapai Co., Verde Canyon, Mexico: Baja California Norte (Sierra San Pedro
[“William A. Tinsley’s Ranch”], G. B. Sudworth s.n. Martír).
(holotype US). TDWG codes: 79 MXN-BC

Bark on trunks smooth and exfoliating in thin

Conservation
flakes and strips, exposing reddish bark, remaining
smooth well into maturity of trees, eventually near In most localities in the Sierra San Pedro Martír
base of large trunks becoming fissured, up to 1 cm where this variety still occurs, the (meta)populations
thick, exfoliating with small flakes. Leaves usually are very small. Fires, but especially grazing by herds
conspicuously and actively glandular. Boss on cone of cattle, are threatening this variety. Even though
scales large or small. Seeds red-brown, not glaucous most known trees are within a National Park, ‘tradi-
or slightly glaucous. tional’ land use like cattle grazing is still continuing
within its boundaries as much as without. It is likely
 that there are more stands than presently known in and the durability of its wood. Changes in fencing
these still largely roadless mountains. techniques on ranches have made this use largely
IUCN: VU (D2) redundant.

Cupressus arizonica Greene var. nevadensis Cupressus arizonica Greene var. stephensonii
(Abrams) Little, Madroño 18: 164. 1966. Cupressus (C. B. Wolf) Little, Madroño 18: 164. 1966.
nevadensis Abrams, Torreya 19: 92. 1919; Callitropsis Cupressus stephensonii C. B. Wolf, Aliso 1: 125. 1948;
nevadensis (Abrams) D. P. Little, Syst. Bot. 31 (3): Callitropsis stephensonii (C. B. Wolf) D. P. Little,
474. 2006 (nom. ut. rej., Art. 56); Hesperocyparis Syst. Bot. 31 (3): 474. 2006 (nom. ut. rej., Art. 56);
nevadensis (Abrams) Bartel, Phytologia 91 (1): 182. Hesperocyparis stephensonii (C. B. Wolf) Bartel,
294 2009. Type: USA: California, Kern Co., Bodfish, Phytologia 91 (1): 183. 2009. Type: USA: California,
Bald Eagle Peak (Clear Creek) [“Red Hill” in Wolf, San Diego Co., King Creek, upper drainage of
1948: 119], L. R. Abrams 5368 (holotype DS). creek, C. B. Wolf RSA 9467 (holotype RSA).

Vernacular names Cupressus arizonica Greene var. revealiana Silba,

Phytologia 49 (4): 393. 1981.
Piute Cypress

Description Vernacular names

Bark on trunks becoming fissured and fibrous, 1–2 Cuyamaca Cypress
cm thick, exfoliating in long strips. Leaves conspic-
uously and actively glandular. Boss on cone scales
Description
usually small and obtuse. Seeds red-brown to dark
brown, more or less glaucous. Bark on trunks smooth and exfoliating in thin
flakes and strips, exposing cherry-red bark, remain-
ing smooth even on large trunks. Leaves variably
Distribution
glandular, from eglandular on some branchlets to
USA: California, in Kern, Los Angeles and Tulare conspicuously and actively on (fewer) others, but
counties, mostly in the Kern River drainage. mostly inconspicuous. Boss on cone scales large or
TDWG codes: 76 CAL small. Seeds red-brown to dark brown, not or more
or less glaucous.
Conservation
Distribution
This variety has a limited distribution in ca. 12 (meta)
populations (groves) of different size in 4 localities, Mexico: Baja California Norte (Sierra Juárez), USA:
the largest being on the N slope of Bald Eagle Peak S California (Cuyamaca Mts.).
S of the little town of Bodfish, covering more than TDWG codes: 76 CAL 79 MXN-BC
200 ha with several thousand trees. These popula-
tions occur both on private and public lands. Fire,
Conservation
cutting or clearing of ‘brush’ to extend pasture, and
competition by shrubs and trees are some of the ma- The two known populations of this variety are very
jor factors that negatively influence the abundance restricted. Fires swept the upper King Creek drain-
of Cypresses. A major burn all but destroyed the age in 1950 and 1970 reducing the size of the popu-
Cannell Creek grove several years ago (J. A. Bartel, lation. A later fire in 2003 further destroyed most
e-mail comm. May 2000). of the remaining mature trees, but regeneration by
IUCN: VU (D2) seed is expected to occur after these events. The
populations in the Sierra Juarez appear to extend as
far as the vicinity of Santa Catarina, with the nearest
Uses
to King Creek ca. 50 km and the farthest ca. 150 km
In the past, trees were cut for fence posts, chosen to the south. Fires are probably less frequent and less
for the small size and straight habit of most trees
severe in Mexico, but there is little or no monitoring itary, subglobose or ovoid, 2–3 × 2 mm, more or less
of the situation. 4-sided; microsporophylls 8–12(–14), decussate, pel-
IUCN: VU (D2) tate, bearing abaxially 2–3(–4) subglobose-angular
pollen sacs. Seed cones terminal, solitary, grouped
but not densely clustered, maturing in two seasons
Cupressus bakeri Jeps., Fl. Calif. 1 (1): 61. 1909. to grey-brown or (silver-)grey subglobose cones of
Callitropsis bakeri (Jeps.) D. P. Little, Syst. Bot. 31 10–18(–22) mm diam. when slightly opened, persist-
(3): 473. 2006 (nom. ut. rej., Art. 56); Hesperocyparis ent. Bract-scale complexes (4–)6–8(–10), decussate,
bakeri (Jeps.) Bartel, Phytologia 91 (1): 180. 2009. peltate, when growing with a prominent boss, when
Type: USA: California, Siskiyou Co., Timbered full grown abaxially rugose, often verrucate, with a
Crater, between Hills Farm (near Dana) and Little near-central, 2–3 mm long, triangular, or obscure, 295
Hot Springs Valley, M. S. Baker s.n. (holotype umbo (bract tip), adaxially constricted to a narrow
JEPS). base, grooved and ridged, dark brown, with lighter
seed scars. Seeds 40–60(–85) per cone, 3–4 × 3 mm,
Cupressus bakeri Jeps. subsp. matthewsii C. B. Wolf, oval to triangular, more or less flattened, brown or
Aliso 1: 83. 1948. red-brown, often glaucous, wings 2, forming thin,
narrow margins up to 0.5 mm wide.
Etymology
Distribution
This species was named after M. S. Baker, who col-
lected the type specimen. USA: N California (Siskiyou Co., Modoc Co., Shasta
Co., Plumas Co.), S Oregon (Josephine Co.).
TDWG codes: 73 ORE 76 CAL
Vernacular names
Baker Cypress, Siskiyou Cypress, Modoc Cypress
Ecology
In Pinus-Quercus woodland and, more commonly,
Description
above that zone in mixed conifer forest or wood-
Trees 10–15(–25) m tall; trunk branching low, to 60– land with Pinus ponderosa, P. jeffreyi, Abies concolor,
100(–140) cm d.b.h. Bark smooth, exfoliating with Pseudotsuga menziesii, Calocedrus decurrens, Arcto-
thin flakes exposing reddish, purplish red or red- staphylos patula, and Seriphidium sp. (Artemisia);
brown bark, giving a shaggy appearance, on lower predominantly on old lava flows in rocky or some-
trunk to 2 cm thick and breaking into small plates times sandy soil. The altitudinal range is between ca.
with upcurled edges, grey. Branches spreading, up- 800 m and 2000 m a.s.l.
per branches ascending, persistent, with foliage to-
wards end, giving a tufted appearance to the often
Conservation
sparse, conical or pyramidal crown. Foliage branch-
es lax or rigid, ultimate branchlets opposite or alter- This species occurs in 9 sites, some quite disjunct;
nate, spreading at 30–60 degrees, variable in length one of these comprises a population more than 3 km
(4–25 mm) but many ca. 10 mm, slender, 0.6–1.2 across, the others are smaller or of unknown size,
mm diam., quadrangular in cross-section, persist- and one population has become extinct in the 20th
ent. Leaves decussate, imbricate, on (sub)ultimate century. Wildfires are the major hazard and dif-
branchlets appressed, rhombic, 1.2–2 × 1 mm, keeled ficult to fight due to the inaccessible terrain of the
with a central depression; margins hyaline-denticu- lava fields in which several of the populations occur.
late or serrulate, obtuse or acute, on whip shoots up More accessible populations are also threatened be-
to 10 × 3–4 mm, often with a spreading apex; sto- cause National Forest fire suppression policy in the
mata few near base abaxially, scattered from base past has allowed the build-up of a dangerous fuel
to apex adaxially; glands conspicuous, central on load, making fires when they occur much more de-
rhombic leaves, producing resin droplets; leaf colour structive.
grey-green to dark green. Pollen cones terminal, sol- IUCN: VU (B1+2bcd)
 shoots with spreading apices) dimorphic (mono-
Uses
morphic on whip shoots), scale-like, with facials
No commercial uses are known of this species; like slightly smaller than laterals, on ultimate branchlets
other cypresses trees may have been felled for fence 1.4–3 × 0.5–1 mm, on whip shoots up to 12 × 2 mm;
posts in the past. Although trees retain a conical laterals conduplicate, curved, their free part spread-
crown which is attractive in gardens, it is little used ing at or below the apex of facials, with incurved or
in horticulture, presumably because it is not strik- more rarely spreading acute apex; facials partly cov-
ingly distinct from other Californian species. ered by laterals, with acute, appressed or free apex;
margins entire; glands conspicuous on facials, in-
conspicuous on laterals; stomata few, scattered pri-
296 Cupressus cashmeriana Royle ex Carrière, Traité marily on margins near leaf bases; leaf colour green
Gén. Conif., ed. 2, 1: 161. 1867. Type: United or glaucous green, sometimes very glaucous. Pollen
Kingdom: [cult. at the Royal Botanic Gardens, Kew cones solitary and terminal on ultimate branchlets,
(“Himalayan House”)], W. J. Bean & [?] Foster s.n. oblong, 4–6 × 2–2.5 mm; microsporophylls 10–16,
(neotype K). decussate, peltate with erose-denticulate margins
and more or less acute apex, bearing 4 abaxial glo-
Cupressus pendula Griff., Itin. Pl. Khasyah Mts.: 131. bose pollen sacs near the lower margin. Seed cones
1848, non Thunb. (1783). solitary or in groups near upper ends of pendulous
Cupressus tortulosa Griff., Not. Pl. Asiat. 4: 26. 1854 branches, occasionally in persistent clusters, termi-
[et in Icon. Pl. Asiat. t. 372. 1854] [“tortulosus” or nal on leafy branchlets with unaltered leaves, ma-
“torulosis”, nom. ut. rej. (proposed 2010)]. turing in 2 growing seasons, (sub)globose to ovoid,
Cupressus himalaica Silba, Phytologia 64: 80. 1987. (10–)12–21 × 10–19 mm., green or glaucous green
maturing to brown with parting scales. Bract-scale
complexes 8–10 in decussate pairs, of more or less
Etymology
equal size except for smaller connate proximal pair,
The species epithet means ‘from Kashmir’ which peltate, 4–5-angular in outline, centrally depressed
was an erroneous conjecture by the author of the with protruding 1 mm long bract tip, rugose, abrupt-
species. [Under the Botanical Code, no name shall ly narrowing towards the cone axis; adaxial surfaces
be changed because it has the wrong meaning.] striated, (red-)brown with light grey seed marks near
base. Seeds 10–15 on each scale (fewer on the proxi-
mal pair), closely packed, angular-ovoid, slightly
Vernacular names
flattened, ca. 4 × 2.5 mm, reddish brown with whit-
Bhutan Cypress, Weeping Cypress ish concave hilum at base, wings 2 on opposite sides,
unequal in size and shape, 1–1.5 mm wide to nearly
absent.
Description
Trees to 85–95 m tall (D. B. Gurung & S. Miehe, un-
Taxonomic notes
publ. data); trunk often buttressed in large speci-
mens, to at least to 3.5 m diam. above the buttress. The taxonomy and nomenclature of this species
Bark becoming fibrous, reddish brown with purplish have been confusing ever since Carrière (1855, 1867)
brown inner bark, exfoliating in long shaggy strips. described two new cypresses with slender, lax foli-
Branches relatively long, spreading or ascending, of- age based on young plants cultivated in the Jardin
ten S-curved, ending drooping to pendulous, form- des Plantes, Paris. Having received these plants from
ing a conical or pyramidal or in old trees irregular another horticulturist without sufficient evidence of
and broad, dense crown. Foliage branches (long) provenance, his statements about origin were specu-
pendulous or more rarely drooping-pendulous, very lative. No original material survives, but of the sev-
slender, lax, ultimate branchlets alternating, disti- eral competing names originally available for the
chous, pendulous, gradually decreasing in length to- Weeping Cypress of Bhutan (C. pendula Griff. 1848
wards tip, together forming planate frondose sprays, unfortunately being a later homonym of C. pendula
ultimate branchlets semi-deciduous. Leaves decus- Thunb. 1783 and C. tortulosa Griff. now proposed
sate, imbricate, decurrent (long decurrent on whip for official rejection for being obscure as well as
a­ mbiguous in its original spelling), C. cashmeriana other traditional buildings, indicate a serious vul-
Carrière could be traced back to 19th century culti- nerability of the remaining wild populations. Pro-
vated plants in Kew that may have the same origin as tection largely takes the form of ex situ conservation
Carrière’s plant (Farjon, 1994). This material is also by planting of individual trees in villages and near
conspecific with Griffith’s collections of C. pendula monasteries and temples.
Griff. and a neotype was therefore selected from it. IUCN: VU (B1+2c)
Cupressus corneyana Carrière remains incertae sedis
and its usage in Flora of Bhutan (Grierson & Long,
Uses
1983) is incorrect. There are apparently very glau-
cous, less glaucous, and non-glaucous forms with The uses of this tree are ornamental, religious, pos-
long, pendulous or shorter, more drooping foliage sibly medicinal; timber harvesting may now have 297
branches. Horticulturists tend to make much of this, ceased largely due to scarcity and inaccessabil-
but we must realise that in almost all cases we are ity of remaining wild populations. This species is
dealing with consciously or unconsciously select- commonly planted in Buddhist monasteries and
ed plants and their mostly clonal offspring. Trees temple grounds in Bhutan (from where it was first
growing in the wild in Bhutan have greener, less described by William Griffith), NE India, Sikkim,
pendulous foliage, and C. pendula Griff. and C. cash- and near Darjiling. It is surely the most ornamen-
meriana Carrière as originally seen and described by tal of the true cypresses and the monks undoubt-
these authors are probably planted selections (culti- edly are responsible for unnamed cultivars with
vars). I see no merit in recognizing the form in na- especially drooping branches and glaucous foliage
ture as a distinct species C. himalaica Silba (e.g. in known to Europeans prior to their discovery of pop-
Grimshaw & Bayton, 2009: 299–300). ulations in the wild, which show a mixture of these
and other characters. Some of these forms are quite
hardy, others only survive in areas with mild, virtu-
Distribution
ally frost-free winters such as in southern Europe.
Indigenous in Bhutan, NE India: Arunachal Pradesh; Probably the most famous tree in Europe is a very
planted widely in the region near Buddhist monas- broadly crowned, multi-stemmed specimen on the
teries and temples in E Nepal, Sikkim, Bhutan, Xi- Isola Madre in Lake Maggiore, northern Italy, now
zang [Tibet], and Arunachal Pradesh. sadly in a poor state after a storm.
TDWG codes: 40 EHM-AP EHM-BH

Cupressus chengiana S. Y. Hu, Taiwania 10: 57. 1964.

Ecology
A very large emergent in evergreen angiosperm for-
Etymology
est dominated by Quercus, with lauraceous trees in
the understorey; also with Tsuga dumosa near the The species epithet commemorates the Chinese bot-
upper limit, and on rocky (limestone) cliffs in pure anist W. C. Cheng.
stands. There are two possible strategies involved:
late successional stands depend on episodal distur-
Vernacular names
bance for regeneration, and extra-zonal avoidance of
competition on exposed cliffs. The altitudinal range min jiang bai mu (Chinese)
is from 1250m to 2670 m a.s.l. The climate in optimal
stands is strongly influenced by summer monsoon
Description
rains, with ca. 800–2000 mm annual precipitation.
Trees to 30 m tall; monopodial; trunk to 1.5 m
d.b.h. Bark eventually becoming fibrous, reddish
Conservation
brown with purplish brown inner bark, exfoliating
The scarcity of localities where this species appears in long strips. Branches spreading, assurgent or as-
to grow naturally, with often few large trees present, cending, often ending drooping, forming a conical
as well as the general desirability of cupressaceous or ­pyramidal or in old trees irregular and broad,
wood in E Asia in connection with religious and dense crown. Foliage branches drooping-­pendulous,
 s­ lender, ultimate branchlets alternating, more or less is extensive, ranging from ca. 1200 m to 2750 m a.s.l.
distichous, (sub)pendulous, forming short feath- The climate is characterized by cold winters and
ery sprays, 1–1.5(–2) mm diam., torose, persistent. cool to warm summers, with a distinct alternation
Leaves decussate, imbricate, decurrent, (nearly) of dry and rainy seasons; annual precipitation varies
monomorphic, scale-like, with facials equal in size between 500–750(–1000 mm), with a 50–70% mois-
to laterals, on ultimate branchlets 1–1.5 mm long and ture deficit. There is no recorded difference in ecol-
rhombic-gibbous, on whip shoots up to 7 mm long, ogy for the two varieties.
with incurved and appressed obtuse apex; margins
minutely erose-hyaline; glands conspicuous on all
Uses
scale leaves; stomata few, scattered primarily on mar-
298 gins near leaf bases; leaf colour dull green or greyish Timber of this species is valued for building and the
green, often covered in thick cuticular wax. Pollen species has been exploited for its larger, more or less
cones solitary and terminal on ultimate branchlets, straight trunks for many centuries. On a small scale
ovoid-oblong, 2–4 × 1–2 mm; microsporophylls trees have been planted as individuals or in groups
12–16, decussate, peltate, bearing 4–5 abaxial pollen in villages.
sacs near the lower margin. Seed cones solitary or in
groups near upper ends of lateral branches, terminal 2 varieties are recognized:
on leafy branchlets up to 10–15 mm long with un-
altered leaves, maturing in 2 growing seasons, per- Cupressus chengiana S. Y. Hu var. chengiana. Type:
sistent, (sub)globose to ovoid-oblong when closed, China: Sichuan, Daxue Shan, Kangding, [“Ta-
(8–)10–25 mm long, maturing to (purplish) brown chien-lu”], W. C. Cheng 2066 (holotype A).
with parting scales. Bract-scale complexes 8–14 in
decussate pairs, of more or less equal size (6–15 mm Cupressus fallax Franco, Portugaliae Acta Biol., sér.
wide), peltate, polygonal (4–5-angular) in outline, B, Sist. 9: 190. 1968.
centrally depressed, flat, or bossed, with protrud-
ing minute bract tip, rugose, abruptly narrowing
Description
towards cone axis; adaxial surfaces striated, brown
with whitish seed marks near base. Seeds 4–6(–8) on Seed cones (sub)globose, very variable in size; bract-
each scale, closely packed, ovoid-triangular, slightly scale complexes 8–14.
flattened, 3–5 × 2–3 mm, reddish brown or yellow-
ish brown with whitish hilum at base; wings 2 on
Distribution
opposite sides, more or less equal in size and shape,
dependent on room for growth 1–2 mm wide. China: S Gansu, N & W Sichuan.
TDWG codes: 36 CHC-SC CHN-SA
Distribution
Conservation
China: S Gansu, N & W Sichuan.
TDWG codes: 36 CHC-SC CHN-GS This variety is known from ca. 9 different locations
where natural populations are now mostly restricted
to more inaccessible sites such as canyons and ex-
Ecology
posed cliffs. Overcutting is the main cause of its
In small, pure stands in some valleys, but more ­decline. A collection from Hunan Province (L. B. Hu
commonly on rocky slopes or cliffs associated with 1233, seen in E) is most likely from an introduced
Koelreuteria paniculata, Morus mongolica, Campy- tree. Cypresses of this species are planted in villages
lotropis delavayi, Bauhinia faberi, Cotoneaster mul- within its natural range in Sichuan and possibly in
tiflorus, and C. gracilis; in non-acidic brown soils Gansu as well.
over granites, quartzites and limestones. Based on IUCN: VU (A1c)
verified herbarium collections, the altitudinal range
Cupressus chengiana S. Y. Hu var. jiangensis ­ ersistent. Leaves covering branchlets decussate,
p
(N. Zhao) Silba, Phytologia 49: 394. 1981. Cupres­ imbricate, decurrent, monomorphic, scale-like, with
sus jiangensis N. Zhao, Acta Phytotax. Sin. 18 (2): facials equal in size to laterals, on ultimate branch-
210. 1980, [“jiangeensis”]. Type: China: Sichuan, lets 1–2 × 0.8 mm (on whip shoots up to 12 mm long,
Longmen Shan, Jiange Xian, L. S. Cai & T. Z. Min often with free apex), rhombic, gibbous or not, with
101–104 (holotype SCFI). incurved and appressed obtuse or acute apex; mar-
gins minutely erose-hyaline; glands conspicuous or
inconspicuous; stomata few, scattered primarily on
Description
margins near leaf bases; leaf colour greyish green
Seed cones ovoid-oblong, more equal in size; bract- or glaucous green; gland sometimes yellowish and
scale complexes 10–12. resinous. Pollen cones solitary and terminal on ul- 299
timate branchlets, obovoid-oblong, (4–)5–7 × 2–3
mm, yellowish green turning light brown; micro-
Distribution
sporophylls (12–)16–20, decussate, peltate, bear-
China: Sichuan (Longmen Shan, Jiange Xian). ing 4–6 abaxial pollen sacs near the lower margin.
TDWG codes: 36 CHC-SC Seed cones solitary or in groups near upper ends of
slender branches, terminal on very short branchlets,
maturing in 2 growing seasons, persistent, globose
Conservation
to angular-globose when open, 15–32 mm diam.,
Only known from a single monoecious tree well to maturing to (purplish) brown with parting scales.
the E of the main populations of var. chengiana. Bract-scale complexes (6–)8–10 in decussate pairs,
IUCN: CR (D) of more or less equal size (up to 15 mm wide) except
for smaller proximal and distal (often connate) pairs,
peltate, polygonal (4–5-angular) in outline; abaxial
Cupressus duclouxiana Hickel, in Camus, [Les face nearly flat, rugose, often pustulate, with grooves
Cyprès] Encycl. Econ. Sylvicult. 2: 91, f. 419–424. radiating from the inconspicuous bract tip in the
1914. Type: China: Yunnan, Kunming Xian, centre, abruptly narrowing towards cone axis; adax-
(introduced), F. Ducloux 3452 (lectotype P). ial surfaces grooved, with whitish seed marks near
the base. Seeds 10 or more around each scale base,
closely packed, ovoid-oblong, slightly flattened, 4–5
Etymology
× 2–3 mm, reddish brown or chestnut-brown with
The species epithet commemorates the French mis- whitish hilum at base; wings 2 on opposite sides,
sionary and plant collector F. Ducloux, who was more or less equal in size and shape, up to 1 mm
based in Kunming. wide.

Vernacular names Distribution

Yunnan Cypress; gan xiang bai (Chinese) China: SW Sichuan, NW Yunnan, SE Xizang [Tibet],
in the deep gorges of the Jinsha (Yangtse), Langcang
(Mekong), and Nu (Salween) Rivers, also along some
Description
western tributaries of the Yalong River in S Sichuan.
Trees to 30(–45) m tall; monopodial; trunk to 1(–1.6) All other references to localities in Yunnan are based
m d.b.h. Bark eventually becoming fibrous, fissured, on introduced trees. The Abbé Delavay understood
reddish brown, several cm thick on lower trunk, this well; he notes with his coll. No. 6805: “grande
slowly exfoliating in long strips. Branches spreading arbre cultivé partout aux environs d’Yunnan-sen.” It
horizontally or ascending in younger trees, form- apparently has established itself successfully in the
ing a conical or pyramidal or in some trees flat- Western Hills S of Kunming and probably elsewhere
topped, dense crown. Foliage branches spreading to in the dry hills of central Yunnan, but that is well
drooping-pendulous, very slender, ultimate branch- outside its natural range. With man’s help (cupres-
lets irregularly disposed, (sub)pendulous, forming saceous trees have been transported all over China
lax, short bushy sprays, 0.8–1 mm diam., torose, and beyond for centuries) the species got out of the
 gorges and onto the presently mild plains and hills extensively in the hills around Kunming, where it
near Lijiang, Dali, and Kunming. ­attains a fastigiate habit quite different from the wild
TDWG codes: 36 CHC-SC CHC-YN CHT growing trees in the great river gorges to the NW. In
Europe it has been introduced in the milder parts of
England, Ireland and France; new provenances from
Ecology
the Kunming trees prove to be hardier and this at-
In mountain forests dominated by Pinus densata tractive species deserves a wider use in horticulture.
and/or deciduous or evergreen Quercus, Castan-
opsis, or Castanea, often gregarious. In many areas
introduced and under deforestation and grazing Cupressus dupreziana A. Camus, Bull. Mus. Hist.
300 pressures invading into scrubland and grassland; Nat. (Paris) 32: 101. 1926.
also widely planted in or near villages. Trees of this
species observed on steep slopes above the Mekong
Etymology
(Langcang) River in NW Yunnan were not associat-
ed with other trees and appeared to grow in the wild. This species was named after “M. le Capitaine Du-
There were trees in all age classes including senes- prez, commandant l’annexe de Djanet” who guided
cent; a grove of very large trees near the river bridge the collector of the type specimen to the trees.
NW of Dêqên is probably a remnant of old growth
forest now protected by the local Buddhist people.
Vernacular names
This species grows naturally in rocky or gravelly
loamy sand or scree over limestone or metamorphic Saharan Cypress, Moroccan cypress; Cyprès de
rocks on the lower slopes of the deep V-shaped river Duprez, Cyprès de l’Atlas (French); tarout (Berber,
valleys under semi-xeric conditions. Its altitudinal ­Tuareg [Tamashek]), azel (Berber?)
range in natural stands is probaly between 1900 m
and 3300 m a.s.l.
Description
Trees to 16–18(–20) m tall; monopodial; trunk to
Conservation
2–3 m d.b.h. Bark eventually becoming thick, deep-
The truly wild and natural (old growth) stands of ly fissured, hard, grey-brown, exfoliating slowly in
this species are extremely rare and scattered in the small strips. Branches long, spreading or ascend-
deep gorges of the great rivers in NW Yunnan, a ing, sometimes fastigiate, forming a conical or py-
few on tributaries. Some are protected by local Bud- ramidal, or in old trees sympodial, irregular and
dhist tradition as holy trees or groves, but adequate broad, dense crown. Foliage branches spreading or
reserves and/or legislation to protect these trees drooping-pendulous, slender, ultimate branchlets
from logging and firewood collecting are absent at irregularly spreading or (sub)pendulous, subterete
present. Regeneration is problematic due to live- to slightly flattened in cross section, 1–1.5(–2) mm
stock grazing, and fire is a potential hazard to the diam., slightly torose, persistent. Leaves covering
few remaining good stands of old growth trees, sev- branchlets decussate, imbricate, decurrent, (nearly)
eral of which are near roads or villages. monomorphic, scale-like, with facials equal in size
IUCN: EN (B1+2de) to laterals, on ultimate branchlets 1–1.2 mm long and
rhombic, slightly gibbous, with appressed obtuse
or acute apex (on whip shoots up to 5 mm long);
Uses
margins minutely denticulate-hyaline; glands on all
Although it is difficult to prove, widespread logging scale leaves (most conspicuous on facials; stomata
may well have depleted forests once more wide- few, scattered primarily on margins near leaf bases;
spread in the river gorges. The wood is similar to leaf colour lustrous greyish green or glaucous green,
that of Taiwania and trees can attain considerable often covered in thick cuticular wax. Pollen cones
height and size. It would produce highly valuable solitary and terminal on ultimate branchlets, ovoid-
timber for local and regional use. This species is also oblong, 4–6 × 2–3 mm; microsporophylls 12–16,
of high horticultural merit and has been planted decussate, peltate, bearing 3–4 abaxial pollen sacs
near the lower margin. Seed cones mostly solitary
Ecology
on ­lateral branches, terminal, maturing in 2 grow-
ing seasons, persistent, subglobose to ovoid-oblong Virtually without any associated vegetation and
when closed, 15–27 × 13–21 mm, maturing to light widely scattered in canyons and dry stream beds
brown with mostly parting scales. Bract-scale com- (wadis) in sandstone plateau; one variety (var. atlan-
plexes (8–)10–12 in decussate pairs, of more or less tica) in very open shrubland or degraded woodland
equal size except for smaller proximal and distal with Juniperus phoenicea, Tetraclinis articulata, and
connate pairs, peltate, polygonal (5–6-angular) in Pistacio lentiscus (in valley bottoms); on rocky lime-
outline; abaxial surface flat, convex or bossed, with stone slopes and in dry, sandy stream beds. The alti-
protruding minute bract tip, (finely) rugose, abrupt- tudinal range of var. dupreziana is 1700–1900 m and
ly narrowing towards cone axis; adaxial surfaces of var. atlantica 900–2220 m a.s.l. 301
grooved, with whitish seed marks near base. Seeds
6–8(–10) on each scale, closely packed, flattened,
Uses
5–6 × 4–5 mm, dark brown with whitish hilum at
base; wings 2 on opposite sides, more or less equal (Illegal) recent use for firewood has been recorded;
in size and shape, 1–2 mm wide. in Morocco also for timber. This species is in cul-
tivation but at present only as ex situ conservation
attempts (mainly concentrated on var. dupreziana
Taxonomic notes
from the Sahara) and probably in a few collec-
The two cypresses indigenous in Algeria and Mo- tions. From observations made on some trees in
rocco have been described seperately as two distinct the wild of both varieties, it should be possible to
species: Cupressus dupreziana A. Camus and C. at- select shoots that grow into a fastigiate habit as with
lantica Gaussen. The only morphological differences C. sempervirens; this could lead to cultivars of inter-
are the shapes of mature seed cones and seeds. How- est to horticulture. Young trees planted in the Royal
ever, even these distinctions are less clear than Gaus- Botanic Gardens, Kew have a columnar shape with
sen (1968) maintained: in C. atlantica they vary from short, more or less ascending branches.
ovoid to subglobose and are rarely if ever globose as
described in the protologue. Gaussen’s description 2 varieties are recognized:
was based on a specimen with a globose cone with
only 8 seed scales, which is somewhat extreme for Cupressus dupreziana A. Camus var. dupreziana.
this taxon. The ultimate branchlets are not thinner Cupressus sempervirens L. var. dupreziana
than in C. dupreziana in most specimens seen. Both (A. Camus) Silba, Phytologia 49: 398. 1981; Tassili­
taxa are also close to C. sempervirens; in fact some cyparis dupreziana (A. Camus) A. V. Bobrov &
specimens seen from western Asia belonging to the Melikyan, Komarovia 4: 72. 2006. Type: Algeria:
natural form of that species (‘horizontalis’) have Oasis, Tassili-n-Ajjir, Tamrit (Wadi), L. Lavauden
similar small cones. However, C. sempervirens does s.n. (holotype P). Fig. 95
not occur naturally west of Crete and Cyrenaica in
Lybia and may have been separated from these taxa
Description
since the early Pleistocene. The disjunction between
the western North African taxa is likely of a more Seed cones ovoid-oblong, with (10–)12 bract-scale
recent date, with forests of Cedrus and Cupressus still complexes; seeds not angular.
existing in the highlands of the Sahara connecting
these with the Atlas Range during the last Ice Age.
Distribution
On these grounds both taxa are here recognized at
infraspecific rank. SE Algeria (Tassili Plateau: Tamrit).
TDWG codes: 20 ALG
Distribution
Conservation
Algeria, Morocco; in Algeria restricted to one very
limited area in the Sahara. This cypress is one of the most extreme examples of a
TDWG codes: 20 ALG MOR-MO relict conifer seemingly on its way out to ­extinction.
 The causes of this are primarily natural climate from ca. 5500 ha in 1950 to 1460 ha in 1986. More
change, but the remaining trees were also cut oc- recent surveys by Griffiths et al. in 1997 (unpubl.
casionally for firewood by wandering Tuareg people exped. rep. Univ. of Reading, UK) failed to observe
and there are tales about caravans that carted off natural regeneration even within an experimental
wood. In all, 233 living individuals are now known enclosure designed to fence out the sheep and goats.
to exist, with 20 trees dead since 1972 also found Exploitation for timber and firewood has certainly
(Abdoun & Beddiaf, 2002). The remaining trees are slowed down under current awareness programmes
often senescent and natural regeneration appears to by the regional authorities, but protection is still in-
be sporadic. The most recent census of trees in the adequate and further decline seems inevitable. Af-
area showed a continuous range of tree sizes and forestation to provide alternatives for local use of
302 found more seedlings, indicating slow but continu- wood is needed, but great care must be taken with
ous natural regeneration. Tourism is providing an the risk of genetic introgression from closely related
increasing incentive for protection and the Tassili species, e.g. C. sempervirens.
n’Ajjer has been designated a World Heritage Site. IUCN: EN (A1bc, B1+2b)
However, adequate protection in situ is still wanting.
Ex situ cultivation is now well established in several
countries. An International Arboretum is being es- Cupressus funebris Endl., Syn. Conif.: 58. 1847.
tablished in Canberra, Australian Capital Territory Chamaecyparis funebris (Endl.) Franco, Agros 24:
(ACT), within which will be established forests of 93. 1941; Platycyparis funebris (Endl.) A. V. Bobrov
rare and endangered species from throughout the & Melikyan, Komarovia 4: 73. 2006. Type: China:
world. One of these forests is dedicated to Cupres- Zhejiang, [“China, prov. of Chekiang, Sir. Geo.
sus dupreziana var. dupreziana and 1300 of the trees Staunton” (introduced)], G. L. Staunton s.n. (lecto­
have been propagated for planting in late 2007. type BM).
IUCN: CR (A2c, C1)
Etymology
Cupressus dupreziana A. Camus var. atlantica
(Gaussen) Silba, J. Int. Conifer Preserv. Soc. 5 (2): The epithet means: ‘of the funeral’ (Latin funus =
29. 1998. Cupressus atlantica Gaussen, Monde Pl. death) and indicates its traditional use in China.
45: 55. 1950; Cupressus sempervirens L. var. atlantica
(Gaussen) Silba, Phytologia 49: 398. 1981. Type:
Vernacular names
Morocco: Haut Atlas, Marrakech, Oued N’Fis, [?]
Idni s.n. (holotype TLF). Funereal Cypress, Chinese Weeping Cypress; bai mu
(Chinese)
Description
Description
Seed cones globose to ovoid-globose, with (8–)10–
12(–11) bract-scale complexes; seeds more or less Trees to 30–35 m tall; monopodial; trunk up to 2 m
angular. d.b.h. Bark eventually becoming shallowly fissured,
grey-brown, exfoliating in long strips. Branches rel-
atively long, spreading or ascending, ending droop-
Distribution
ing to pendulous, forming a conical or pyramidal
Morocco (Oued-n’Fis valley). or in old trees irregular and broad crown. Foliage
TDWG codes: 20 MOR-MO branches pendulous, slender, lax, ultimate branch-
lets distichous, pendulous, forming planate fron-
dose sprays, slightly flattened (subtetragonous) in
Conservation
cross section, ultimate branchlets semi-deciduous.
The main population occurs in the Oued-n’Fiss val- Leaves covering decussate, imbricate, decurrent, di-
ley between Asni and Ijoukak, where many old trees morphic (monomorphic on whip shoots), scale-like,
still survive. Estimates indicate however that the to- with facials slightly smaller than laterals, on ultimate
tal area of occupancy (AOO) suffered a reduction branchlets 1.5–3.5 × 0.5–1 mm, on whip shoots up to
10 × 2 mm; laterals conduplicate, straight or curved,
Ecology
with appressed or more rarely free acute apex; ­facials
partly covered by laterals, with acute, appressed In mixed mountain forest or (degraded) woodland
apex; margins entire; glands on facials and laterals associated with Platycarya strobilacea, Vitex negun-
(often more conspicuous on facials), linear; stomata do, Ligustrum sp., Viburnum sp., Pittosporum sp.,
few, scattered primarily on margins near leaf bases; Myrsine africana, and Vitex negundo; in calcareous
leaf colour lustrous green. Pollen cones solitary soil or in sandy loam over sandstone; also widely
and terminal on ultimate branchlets, ovoid-oblong, planted and probably invading into disturbed veg-
3–5 × 2–2.5 mm; microsporophylls 10–14, decus- etation locally. The altitudinal range of this species is
sate, peltate, bearing 4 abaxial pollen sacs near the between 300 m and 2260 m a.s.l.
lower margin. Seed cones solitary or in groups on 303
pendulous branches, terminal, maturing in 2 grow-
Conservation
ing seasons, caducous, (sub)globose when closed,
(8–)10–15 mm diam., maturing to dark brown with The wide distribution of this species cited in the Chi-
parting scales. Bract-scale complexes 6–8(–12) in nese literature (including Flora of China 4: 67. 1999)
decussate pairs, of more or less equal size except the is to a large extent based on planted or introduced
smaller connate proximal pair, (5–10 mm wide), pel- trees outside the indisputably wild populations. Oc-
tate, polygonal (4–5(–6)-angular) in outline; abaxial currence in natural forests is rare due to widespread
face more or less umbilicate, with a central curved deforestation and alteration of natural vegetation in
boss surrounding the protruding bract tip, finely ru- much of central China. The possibility of secondary
gose, abruptly narrowing towards cone axis; adax- establishment from cultivated trees outside its origi-
ial surfaces angular, striated, chestnut-brown with nal range makes evaluation of threat to wild popula-
whitish seed marks at base. Seeds 3–5(–6) on each tions of this species extremely difficult. This species
scale, closely packed, angular-ovoid, slightly flat- is not considered to be threatened, but its natural
tened, 3 × 2.5 mm, dark brown with whitish hilum at habitat (mixed conifer-angiosperm forest) certainly
base; wings 2 on opposite sides, unequal in size and is. It should perhaps be given the Near Threatened
shape, depending on room for growth 1 mm wide to (NT) category.
nearly absent on one side. IUCN: LC

Taxonomic notes Uses

This species has often been classified in the genus This species is widely planted in China as an or-
Chamaecyparis on the basis of its flattened foli- namental, especially common in Buddhist temple
age (dimorphic facial and lateral leaves on foliage grounds. Its wood is also considered valuable due to
sprays) and small cones. The ontogeny of its seed its durability and traditionally it was used for coffins
cones, which mature in two years, and the higher for this reason. In Europe it is uncommon in culti-
number of ovules/seeds, which arise in two succes- vation and most of the older extant trees are based
sive rows axillary to each bract, place it in Cupressus. on seed collection by Ernest Wilson made in Hubei
The dimorphic leaves are also found in other taxa Province, China, in 1907. In horticulture it is often
in the family adapted to higher athmospheric mois- referred to as Chamaecyparis funebris due to the
ture, and are likely the result of convergent evolu- flattened foliage and small cones.
tion.

Cupressus goveniana Gordon, J. Hort. Soc. London

Distribution
4: 295. 1849.
China: S Gansu (?), Guizhou, W Hubei (?), W Hu-
nan, Shaanxi (?), Chongqing; widely cultivated in
Etymology
Central and S China.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- This species was named after James R. Gowen, at the
SC CHS-HN time Secretary of the Horticultural Society of Lon-
don.
 Vernacular names Distribution
Gowen Cypress, Mendocino Cypress, North Coast USA: California, Mendocino, Sonoma, Santa Cruz,
Cypress, Pygmy Cypress, Dwarf Cypress, Santa San Mateo and Monterey Counties.
Cruz Cypress TDWG codes: 76 CAL

Description Ecology
(Dwarf) trees to 10(–50) m tall; monopodial; trunk In chaparral, ‘pine barrens’, and open pine woodland
to 1(–2.4) m d.b.h. Bark eventually becoming fibrous, with Pinus attenuata, P. contorta, P. muricata, P. pon-
304 greyish brown to grey, hard, fissured, with anasto- derosa, P. radiata, Pseudotsuga menziesii, Arctosta-
mosing ridges, exfoliating in shreddy strips, to 2–3 phylos, Quercus, and Rhododendron, often in groves
cm thick. Branches ascending or spreading, short of up to 1000 trees or more; on sandstone outcrops,
or long, slender, often sparse, forming a conical or white or yellow sandy slopes, and leached, virtually
pyramidal crown, or in certain conditions dwarfed. sterile sandy ‘hardpan’, where it becomes dwarfed.
Foliage branches spreading, slender, ultimate branch- The altitudinal range is from near sea level to 1200
lets irregularly disposed, spreading, short (3–5 mm) m a.s.l. The climate is of the Mediterranean type
or longer (10–20 mm) and rigid forming dense tufts, with dry, hot summers, but in a narrow coastal strip
subterete in cross section, 1–1.5 mm diam., torose, cooled by frequent fog, and winter rain.
persistent. Leaves covering branchlets decussate,
imbricate, decurrent on whip shoots and then with
Uses
recurved, free distal parts, on ultimate branchlets
appressed, monomorphic, scale-like, with facials Although introduced by C. T. Hartweg to England
equal in size to laterals, on ultimate branchlets 1–1.5 in 1848, this species soon turned out to be tender
× 1–1.3 mm and rhombic-gibbous, on whip shoots up in NW Europe and its cultivation outside collec-
to 5 mm long, with incurved and appressed obtuse tions ceased. In southern Europe it is grown more
or more or less apiculate apex, sometimes slightly widely in gardens and parks and a few cultivars are
keeled; margins minutely denticulate; glands incon- known, some with doubtful affinity to this species
spicuous; stomata few, scattered primarily on mar- (see Vidaković, 1991: 193).
gins near leaf bases; leaf colour yellowish green or
dark green. Pollen cones numerous, solitary and 2 varieties are recognized:
terminal on ultimate branchlets, ovoid-oblong, 3–4
× 1.5–2 mm; microsporophylls 12–14, decussate, pel- Cupressus goveniana Gordon var. goveniana. Calli­
tate, bearing 3–5(–6) abaxial pollen sacs near lower tropsis goveniana (Gordon) D. P. Little, Syst. Bot.
margin. Seed cones in groups or clusters on lateral 31 (3): 473. 2006, (nom. ut. rej., Art. 56); Hespero­
branches, terminal, maturing in 2 growing seasons, cyparis goveniana (Gordon) Bartel, Phytologia 91
persistent, (sub)globose when closed, 13–30 × 12–25 (1): 181. 2009. Type: USA: California, Monterey Co.,
mm, maturing to lustrous brown and finally grey- Huckleberry Hill, [“discovered by Mr. Hartweg on
brown or grey with parting scales. Bract-scale com- the western declivity of the mountains of Monterey
plexes 6–10, usually 8, in decussate pairs, of more in Upper California”], C. T. Hartweg s.n. (holotype K).
or less equal size, 8–15(–18) mm wide, peltate, po-
lygonal (4–6-angular) in outline, nearly flat or more Cupressus goveniana Gordon var. pigmaea Lem-
often bossed, with protruding bract tip, abruptly mon, Handb. W. Amer. Cone-bearers, ed. 3: 77.
narrowing towards cone axis; abaxial surface rugose; 1895; Cupressus pigmaea (Lemmon) Sarg., Bot. Gaz.
adaxial surfaces striated, brown or red-brown with (Crawfordsville) 31: 239. 1901; Cupressus goveniana
whitish seed marks near base. Seeds 8–10 on each Gordon subsp. pigmaea (Lemmon) A. Camus, [Les
scale, closely packed, usually angular, slightly flat- Cyprès] Encycl. Econ. Sylvicult. 2: 50. 1914, [“pyg-
tened, 3–5 × 2–4 mm, (blackish) brown or glaucous maea”]; Callitropsis pigmaea (Lemmon) D. P. Little,
brown with whitish hilum at base; wings 2 on oppo- Syst. Bot. 31 (3): 474. 2006, (nom. ut. rej., Art. 56);
site sides, unequal in size and shape or discontinu- Hesperocyparis pigmaea (Lemmon) Bartel, Phytolo-
ous, depending on room for growth to 1 mm wide. gia 91 (1): 182. 2009 (“pygmaea”).
 the Bonny Doon Ecological Reserve, several others
Description
remain unprotected.
Seed cones 13–20 mm long; seeds (blackish) brown, IUCN: EN (C2a)
not or only slightly glaucous.

Cupressus guadalupensis S. Watson, Proc. Amer.

Distribution
Acad. Arts 14: 300. 1879.
USA: California, Mendocino, Sonoma and Monterey
Counties.
Etymology
TDWG codes: 76 CAL
The species epithet refers to Guadalupe Island, where 305
this species (var. guadalupensis) is indigenous.
Conservation
Only known from a few scattered locations in a lim-
Vernacular names
ited area, some of which are very close to continuing
urban development. Guadalupe Cypress, Tecate Cypress, Forbes Cypress;
IUCN: VU (D2) ciprés (Spanish)

Cupressus goveniana Gordon var. abramsiana

Description
(C. B. Wolf) Little, Phytologia 20: 435. 1970.
Cupressus abramsiana C. B. Wolf, Aliso 1: 215. 1948; Large shrubs or small trees 10–15 m tall, rarely to
Callitropsis abramsiana (C. B. Wolf) D. P. Little, 20 m; trunk short, branching with several lead-
Syst. Bot. 31 (3): 473. 2006, (nom. ut. rej., Art. 56); ers low above ground, or sometimes monopodial
Hesperocyparis abramsiana (C. B. Wolf) Bartel, with an unbranched trunk 2–3 m tall, to 1.2–1.5 m
Phytologia 91 (1): 180. 2009. Type: USA: California, diam. Bark remaining smooth and thin (to ca. 1 cm)
Santa Cruz Co., Bonnie Doon, ca. 1 km E of public even on lower part of large trunks, exfoliating in
School, C. B. Wolf RSA 6235 (holotype RSA). small flakes, or occasionally in long strips remain-
ing partly attached, exposing reddish or purplish
brown to light chocolate-brown bark; outer bark
Description
weathering whitish grey. Branches numerous, long,
Seed cones (15–)20–30 mm long; seeds brown, often thick, spreading, ascending or erect, forming in free
glaucous. standing trees with short trunks a wide and round-
ed, but in more crowded conditions a narrower or
irregular crown. Foliage branches lax or rigid, ulti-
Distribution
mate branchlets 3–15 mm long, slender or thicker,
USA: California, San Mateo and Santa Cruz Counties. 1.2–1.7 mm diam., quadrangular in cross-section,
TDWG codes: 76 CAL becoming thicker and subterete on older branch-
lets, covered with leaves. Leaves decussate, imbri-
cate, on ultimate branchlets appressed, rhombic,
Conservation
1–1.5 × 1 mm, gibbous or with a slight central de-
Restricted to four populations in the Santa Cruz pression; margins ­hyaline-denticulate; apex obtuse,
Mountains; two were located or found since the field up to 15 × 3–4 mm on some vigorous whip shoots;
studies by Wolf (in Wolf & Wagener, 1948). Some, stomata on abaxial side scattered in lower part, on
like the one near Eagle Rock, consist of only a few the adaxial side from base to apex; leaves eglandu-
score mature trees surrounded by highly flamma- lar (ultimate branchlets) or with an inconspicuous,
ble chaparral, and/or are inaccessible to fire fighters inactive gland, covered with a thick cuticular wax
other than from the air. The population near Bonnie layer, glaucous green to light green. Pollen cones
Doon is partly in an area with private real estate de- numerous, ­terminal, solitary, ovoid-oblong, terete
velopment and fire prevention here will allow pines or quadrangular in cross-section, 4–7 × 2–3 mm;
like Pinus attenuata and P. ponderosa to crowd out microsporophylls 12–18(–20), decussate, imbricate,
the cypresses. The main population here is within peltate-cordate, abaxially bearing 3–4(–5) angular-
 globose, yellow pollen sacs. Seed cones terminal, not Cupressus guadalupensis S. Watson var. guada-
aggregated in dense clusters but often grouped with lupensis. Callitropsis guadalupensis (S. Watson)
up to 20 cones, growing in two seasons to globose D. P. Little, Syst. Bot. 31 (3): 473. 2006, (nom. ut. rej.,
or subglobose cones, 15–35 mm diam. (some sub- Art. 56); Hesperocyparis guadalupensis (S. Watson)
globose cones to 45 mm long), serotinous, coarsely Bartel, Phytologia 91 (1): 181. 2009. Type: Mexico:
rugose, sometimes with prominent bosses on the Baja California Norte, Guadalupe Island, E. J. Palm-
scales. Bract-scale complexes (6–)8–10, decussate, er 92 (lectotype K).
peltate, 4–5 sided or partially more or less rounded in
outline, abaxially with a prominent and upcurved or
Description
flat and inconspicuous boss near the centre, coarsely
306 rugose to verrucose, adaxially abruptly constricted, Pollen cones 5–7 mm long, before dehiscence dis-
dark brown with light seed scars towards base. Seeds tinctly quadrangular in cross-section, microsporo-
up to 100–120 per cone, irregularly shaped, slightly phylls 16–18(–20) per cone. Seed cones up to 45 mm
flattened, 4–7 × 2–3.5 mm, dark brown, the hilum long but size highly variable.
conspicuously lighter; wings 2 narrow, irregular
strips on either side, 0.5–1 mm wide.
Distribution
Mexico: Baja California Norte, restricted to Guada-
Distribution
lupe Island.
Mexico: Baja California Norte, Guadalupe Island, TDWG codes: 79 MXI-GU
also along the border with California; USA: SW
California, a few localities in Orange Co. and San
Conservation
Diego Co.
TDWG codes: 76 CAL 79 MXI-GU MXN-BC The situation with the variety of C. guadalupensis
that is restricted to the island of Guadalupe in the
Pacific Ocean is extremely critical due to overgraz-
Ecology
ing by feral goats. The population may not exceed
In chaparral on slopes with Adenostoma spp., Arc- 200 mature trees in two small, separate stands. Re-
tostaphylos sp., in ravines in the Upper Sonoran Life cent fires have destroyed many trees. No permanent
Zone associated with Acer sp., Rhus laurina, Quercus residents live on the island, which is visited only oc-
spp., and Arctostaphylos sp., also locally associated casionally.
with Pinus radiata var. binata; often along intermit- IUCN: CR (B1+2c)
tent streams on loamy, sandy, gravelly or rocky soils
(or ‘adobe soil’) over sandstone or granite in full sun. Cupressus guadalupensis S. Watson var. forbesii
The altitudinal range of var. guadalupensis is from (Jeps.) Little, Phytologia 20: 435. 1970. Cupressus
1040 m to 1230 m and of var. forbesii from 210 m forbesii Jeps., Madroño 1: 75. 1922; Cupressus
to 1400 m a.s.l. The climate is of the Mediterranean guadalupensis S. Watson subsp. forbesii (Jeps.)
type with dry, hot summers and winter rain; with Beauch., Aliso 9: 191. 1978; Callitropsis forbesii
frequent fog on Guadalupe Island. (Jeps.) D. P. Little, Syst. Bot. 31 (3): 473. 2006,
(nom. ut. rej., Art. 56); Hesperocyparis forbesii
(Jeps.) Bartel, Phytologia 91 (1): 181. 2009. Type:
Uses
USA: California, San Diego Co., Otay Mt.,
No uses are known of this species and its varieties. [“Cedar Canyon, between El Nido and Dulzura”],
Only a few botanical collections in California (e.g. C. N. Forbes s.n. (lectotype JEPS). Fig. 96
Rancho Santa Ana Botanical Garden) have grown
this species successfully. It should be taken into cul-
Description
tivation more widely especially for reasons of ex situ
conservation of the variety native to Guadalupe Is- Pollen cones 4–5 mm long, usually terete or some-
land. times slightly angular; microsporophylls 12–14 per
cone. Seed cones globose, 15–30 mm diam.
2 varieties are recognized:
 branches numerous, spreading or drooping, slender,
Distribution
ultimate branchlets irregularly disposed or more or
Mexico: Baja California Norte (near US border), less distichous, ± quadrangular in cross section or
USA: SW California, one locality in Orange Co. and slightly flattened, 1–2 mm diam., persistent. Leaves
a few in San Diego Co. down to the Mexican border. covering branchlets, decussate, imbricate, decur-
TDWG codes: 76 CAL 79 MXN-BC rent, monomorphic to slightly dimorphic, scale-like;
facials equal or somewhat unequal in size to laterals,
on ultimate branchlets 1–2.5 mm long and rhombic,
Conservation
on whip shoots up to 10 mm long; margins minutely
Tecate Cypress has a restricted distribution in San erose-hyaline; apex incurved and appressed acute
Diego County and one disjunct population in Or- (or obtuse); laterals similar or conduplicate, with 307
ange County. The largest population is on Otay appressed or free apex; glands inconspicuous or ab-
Mountain close to the Mexican border; individuals sent esp. on laterals; stomata few, scattered prima-
are known from the Mexican side as well. These lat- rily on margins near leaf bases; leaf colour green or
ter trees are (were) very close to the highway from glaucous green. Pollen cones solitary and terminal
Tecate to Mexicali. Urbanisation in the region has on ultimate branchlets, slightly oblong, more or less
brought an increased risk of wildfires and the pre- quadrangular, 3–5 ± 2–2.5 mm; microsporophylls
vention as well as the attempts to put these down 10–16(–18), decussate, peltate, bearing 3–4 abaxial
or restrict them will be concentrated around urban pollen sacs near lower margin. Seed cones solitary
properties, not in the first place populations of rare or in groups near the upper ends of lateral branches,
trees. Like its congeners in California, this cypress terminal, maturing in 2 growing seasons, persistent,
will regenerate after fire but there is a definite risk to (sub)globose-angular when closed, 10–18(–20) mm
survival if frequency or intensity of fires are increas- diam., maturing to brown with parting scales. Bract-
ing due to human impact factors. scale complexes 6–8(–10) in decussate pairs, of more
IUCN: VU (D2) or less equal size, peltate, polygonal (4–6-angular) in
outline, bossed, with protruding recurved bract tip
3–5 mm long, abruptly narrowing towards the cone
Cupressus lusitanica Mill., Gard. Dict., ed. 8: axis, rugose abaxially; adaxial surfaces striated, with
Cupressus No. 3. 1768. whitish seed marks near base. Seeds 8–12 on each
scale, closely packed, slightly flattened, 3–4.5 × 3–4
mm, brown or yellowish brown with whitish hilum
Etymology
at base; wings 2 on opposite sides, sometimes ab-
The species epithet derives from the Latin name for sent, depending on room for growth 1–1.5 mm wide.
Portugal, Lusitania, where it was first noticed by
botanists.
Taxonomic notes
See the Monograph of Cupressaceae and Sciadopitys
Vernacular names
(Farjon, 2005a) for a discussion of the identity and
Mexican Cypress, Cedar of Goa; Cedro blanco, Ce- provenance of the ‘Bussaco cypresses’ in Portugal
dro blanco del deserto, Cedro, Ciprés (Spanish) (Cupressus lusitanica Mill.) and a comprehensive ci-
tation of the pertinent literature.
Description
Distribution
Trees to 30–35 m tall, monopodial, large trees but-
tressed; trunk up to 2 m d.b.h. Bark eventually be- Mexico: from the Sierra Madre Occidental and
coming fissured and fibrous, brown weathering to S. M. Oriental down to Chiapas; highlands of Guate-
dark grey-brown, exfoliating in long strips. Branches mala, Belize, Honduras, El Salvador and Nicaragua.
long, spreading or ascending, often ending droop- TDWG codes: 79 MXC-DF MXC-ME MXC-PU MXE-
ing-pendulous, forming a pyramidal or in old trees AG MXE-CO MXE-CU MXE-DU MXE-HI MXE-QU
sympodial, rounded or flat-topped crown. Foliage MXE-SL MXG-VC MXN-SI MXS MXT-CI 80 GUA HON
 Ecology Description
Forming pure, dense stands or scattered in mixed Trees, when mature, developing broad crowns. Foli-
montane conifer forest or pine forest, also in pine- age branches not in more or less planate sprays but
oak forest and woodland, associated with Abies spp., irregularly arranged; ultimate branchlets irregularly
Pinus ayacahuite, P. hartwegii, P. maximinoi, P. mon- decreasing in length, more or less quadrangular in
tezumae, P. patula, P. pseudostrobus, Pseudotsuga cross section. Leaves monomorphic, apices of facials
menziesii var. glauca, Juniperus spp., Quercus spp., appressed, of laterals often free; glands present on all
Alnus spp., Clethra sp., Persea sp. and ericaceous scale leaves.
and theaceous undershrubs; in disturbed (grazed)
308 woodland with Arbutus sp., Baccharis sp., Buddleia
Distribution
sp., and Leucena sp. This species occurs on various
usually nutrient-poor rocky soils over limestone or Mexico: from the Sierra Madre Occidental and
various igneous rocks; it is also spreading in scrub S. M. Oriental down to Chiapas; highlands of Guate-
on rocky slopes or cliffs in canyons. The altitudinal mala, Belize, Honduras, El Salvador and Nicaragua.
range of this species is from ca. 1000 m to nearly TDWG codes: 79 MXC-DF MXC-ME MXC-PU MXE-
4000 m a.s.l. AG MXE-CO MXE-CU MXE-DU MXE-HI MXE-QU
MXE-SL MXG-VC MXN-SI MXS-CL MXS-GR MXS-JA
MXS-MI MXS-NA MXS-OA MXT-CI 80 GUA HON
Uses
Mexican cypress is a valuable timber tree where
Conservation
it grows tall and straight, but not used extensively
in its native range. It is planted as a ‘village tree’ IUCN: LC
throughout Mexico and Guatemala; it has also been
introduced in many countries, especially in Andean Cupressus lusitanica Mill. var. benthamii (Endl.)
South America and E Africa and to a limited extent Carrière, Traité Gén. Conif., ed. 2, 1: 155. 1867.
in SE Asia, as a plantation forest tree. Still known in Cupressus benthamii Endl., Syn. Conif.: 59. 1847;
some quarters as the ‘Cedar of Goa’ its true prov- Callitropsis benthamii (Endl.) D. P. Little, Syst.
enance was long a mystery. The Portuguese had a Bot. 31 (3): 473. 2006, (nom. ut. rej., Art. 56);
colony of that name on the west coast of India and Hesperocyparis benthamii (Endl.) Bartel, Phytologia
the species was first described from Portugal but lat- 91 (1): 181. 2009. Type: Mexico: Hidalgo, El Banco,
er understood not to be native there: 1 + 1 = 2 seems [“from the Banco”], C. T. Hartweg 434 (holotype
to have been the reasoning. In horticulture sev- BM). Fig. 97
eral cultivars are known, while var. benthamii, with
more flattened foliage sprays, is also being planted
Description
in countries with mild winters. The latter produces a
narrowly conical habit in cultivation. Trees usually with conical crowns. Foliage branch-
es forming more or less planate sprays; ultimate
2 varieties are recognized: branchlets more or less distichous, gradually de-
creasing in length, slightly flattened. Leaves slightly
Cupressus lusitanica Mill. var. lusitanica. Callitrop- dimorphic; apices of facials and laterals appressed;
sis lusitanica (Mill.) D. P. Little, Syst. Bot. 31 (3): 474. laterals mostly eglandular.
2006, (nom. ut. rej., Art. 56); Hesperocyparis lusita­
nica (Mill.) Bartel, Phytologia 91 (1): 181. 2009. Type:
Distribution
Portugal: Bussaco, [type locality (introduced)], leg.
ign. [ex herb. Ph. Miller] s.n. (holotype BM). Mexico: Chiapas, Guerrero, Hidalgo, México, Pue-
bla, Veracruz; probably scattered in forests with var.
Cupressus lindleyi Klotzsch ex Endl., Syn. Conif.: 59. lusitanica elsewhere.
1847. TDWG codes: 79 MXC-ME MXC-PU MXE-HI MXG-
Cupressus lusitanica Mill. var. hondurensis Silba, VC MXS-GR MXT-CI
Phytologia 68: 30. 1990.
 Pollen cones numerous, terminal, solitary, angular-
Conservation
ovoid, 3–4 × 2–2.5 mm; microsporophylls (8–)10–12,
IUCN: NT decussate, more or less flat or slightly convex, on the
abaxial lower side bearing (3–)4–6(–7) yellow pollen
sacs. Seed cones terminal, often clustered and ap-
Cupressus macnabiana A. Murray bis, Edinburgh pearing sessile, in two growing seasons maturing to
New Philos. J., n.s., 1: 293. 1855, [“M’Nabiana”]. dark brown, eventually grey, subglobose, serotinous
Callitropsis macnabiana (Hartw.) D. P. Little, and persistent cones 15–25 × 13–20 mm. Bract-scale
Syst. Bot. 31 (3): 474. 2006, (nom. ut. rej., Art. 56); complexes (4–)6(–8), decussate, connate, parting
Hesperocyparis macnabiana (A. Murray bis) Bartel, very slowly, peltate, irregularly 4–5(–6) sided, abaxi-
Phytologia 91 (1): 182. 2009. Type: USA: California, ally with a subapical, broadly based, upcurved boss 309
[“Habitat in California, circa lat. 41 Bor.”], 3–6 mm long, terminating in an acute bract tip 1 × 2
W. Murray & A. F. W. Beardsley s.n. (holotype E). mm; outer surface rugose; inner part abruptly con-
Pl. 12 stricted, angular, dark brown, with lighter coloured
seed scars. Seeds 70–100 per cone, 3–4(–5) × 2–3
mm, irregularly shaped, slightly flattened, some-
Etymology
times more or less verrucose, brown, with a lighter
This species was named after William McNab (1780– 1–1.5 mm long hilum; wings 2, forming very narrow
1848) a Scottish botanist. strips around the seed.

Vernacular names Distribution

MacNab Cypress, Shasta Cypress, Fragrant Cypress USA: California, Coast Ranges, mountains of N Cal-
ifornia, Sierra Nevada.
TDWG codes: 76 CAL
Description
Large shrubs to small trees to 10–15 m tall; trunk
Ecology
short, branching very low or from ca. 1 m above
ground, to 80–100 cm diam. Bark on trunk to 3 cm Cupressus macnabiana occurs in the Upper Sonoran
thick, fissured, hard, very slowly exfoliating, eventu- Life Zone in chaparral or woodland, associated with
ally more or less fibrous, grey. Branches numerous, Pinus attenuata, P. sabiniana, less commonly P. pon-
long, slender, spreading, assurgent to erect, foliage derosa, Quercus spp., and Arctostaphylos sp., often
with long, assurgent whip shoots, forming a spread- forming groves on rocky slopes and in ravines in
ing, round crown with branches near the ground. clay, loam or sand over serpentine. The altitudinal
Foliage branches spreading or assurgent, the high- range is from 300 m to 1200 m a.s.l. The climate is of
est order branchlets more or less in one plane but the Mediterranean type with dry, hot summers and
not plagiotropic, mostly ascending, rigid, ultimate winter rain.
branchlets fine, spreading, 3–7(–10) × ca. 1 mm,
torulose, on thicker, older branchlets rough with
Conservation
numerous spreading leaf tips, covered with leaves,
persistent. Leaves decussate, slightly imbricate, ap- IUCN: LC
pressed on smallest branchlets, rhombic to oblan-
ceolate, rounded or keeled abaxially, 1–1.5(–2) ×
Uses
1 mm, with a near central depression or subapical
pit; margins remotely denticulate; apex obtuse; on This relatively small tree may have been locally used
thicker branches and whip shoots to 12 × 3 mm, line- for fence posts by ranchers; today its wood is not
ar-lanceolate, with a reflexed, apiculate or acute dis- considered to be of any commercial value. It is rare
tal part to 3 mm long; stomata near base abaxially, in horticultural cultivation but would be suitable as
scattered from base to apex adaxially; gland mostly an amenity tree in dry climate. Cultivars are scarcely
present and conspicuous, active; leaf colour grey- known; Vidaković (1991) listed ‘Sulphurea’ with yel-
green to green, new foliage sometimes yellow-green. low-tipped foliage branches.
310

5
2
5
1
1

Pl ate 12 . Cupressus macnabiana. 1. Branch with foliage and seed cones. 2. Branchlet with leaves and
pollen cones. 3. Leaves with gland. 4. Microsporophylls with pollen sacs and pollen. 5. Seeds.
Cupressus macrocarpa Hartw. ex Gordon, J. Hort. clustered and appearing sessile, in two growing
Soc. London 4: 296. 1849. Callitropsis macrocarpa seasons maturing to dark brown, eventually grey-
(Hartw.) D. P. Little, Syst. Bot. 31 (3): 474. 2006, brown, subglobose to broadly ovoid, serotinous and
(nom. ut. rej., Art. 56); Hesperocyparis macrocarpa persistent cones 20–35(–40) × 18–30 mm. Bract-
(Hartw. ex Gordon) Bartel, Phytologia 91 (1): scale complexes 8–12(–14), decussate, connate, part-
182. 2009. Type: USA: California, Monterey Co., ing very slowly, peltate, of unequal size (but pairs
[“near Carmel Bay, Monterey”], C. T. Hartweg 143 usually similar) irregularly 4–5(–6) sided or with
(holotype K). Fig. 98 more or less rounded upper margin, abaxially with
a relatively small, broadly based, upcurved boss 2–4
mm long, terminating in an acute bract tip 1 × 2
Etymology
mm, eroding away in old cones; outer surface ru- 311
The species epithet (Latin macro = large; carpos = gose; inner part abruptly constricted, angular, dark
fruit) refers to the relatively large seed cones. brown to blackish, with conspicuous, white seed
scars. Seeds 100–150 per cone, 3–5(–6) × 2–4 mm,
slightly flattened, angular, brown to blackish brown,
Vernacular names
with a whitish hilum; wings 2, forming narrow, ir-
Monterey Cypress, Ciprés Monterrey, Ciprés de Cal- regular strips ca. 1 mm wide.
ifornia (Spanish)
Distribution
Description
USA: California (near Monterey).
Trees to 20–25 m tall; trunk straight, monopodial TDWG codes: 76 CAL
in sheltered places, branching very low from less
than 1 m above ground in exposed places, to 150–170
Ecology
cm diam. Bark on trunk to 4–5 cm thick, fissured,
hard, with anastomosing ridges, very slowly exfolia- Restricted to a narrow coastal strip on rocky cliffs,
ting, grey or ash-grey. Branches long, spreading or slopes and headlands, forming pure stands or as-
ascending, forming a conical or pyramidal crown, sociated with Pinus radiata, in loam or sand over
in old, exposed trees very dense, forming horizon- granitic rocks or in rock crevices. The climate is of
tal canopies and a broad, flat-topped crown. Foli- the Mediterranean type, with dry summers cooled
age branches in tapering or flattened foliage sprays, by frequent fog, within reach of ocean salt sprays,
the highest order branchlets irregularly disposed at and winter rain.
various angles and mostly ascending, rigid, ultimate
branchlets 3–10(–15) × 1.2–1.8(–2) mm, quadrangu-
Conservation
lar in cross-section, covered with leaves, persistent.
Leaves decussate, imbricate, appressed on smallest The limited distribution of this species in its natural
branchlets, on ultimate and penultimate branchlets habitat consists of two main ‘groves’ and a few scat-
(broadly) rhombic, 1–2 × 1 mm, rounded or keeled tered metapopulations. These are mostly situated in
abaxially, with a near central or two lateral depres- reserves (Del Monte Forest, Point Lobos), but also
sions; margins hyaline-denticulate; apex obtuse; on private lands in the vicinity of ongoing urban and
leaves on whip shoots to 20 × 4 mm, long decurrent, leisure development. As recreation and tourism are
linear-lanceolate, with a reflexed, apiculate to pun- intensive in the area, there is a great risk from fires.
gent distal part to 5 mm long; stomata scattered or IUCN: VU (D2)
in two short bands to below apex abaxially, scattered
from base to apex adaxially; glands absent or in-
Uses
conspicuous and inactive; leaf colour green to dark
green. Pollen cones terminal, solitary, angular-ovoid This species of cypress has been widely introduced
to oblong, 3–6 × 2.5–3 mm; microsporophylls 12–14, in California and around the world, in some African
decussate, more or less flat or slightly convex, on the countries as a timber tree, but mostly as an orna-
abaxial lower side bearing 6–8(–10) crowded, ob- mental tree and to act as wind shelter belts both in
long, yellow pollen sacs. Seed cones ­terminal, ­often argiculture and horticulture. The wood from closed-
 grown trees in plantations can be used for carpentry apiculate or pungent apex; margins hyaline-den-
and furniture. The tree is extremely resistant to wind ticulate; stomata on abaxial side restricted to base,
and tolerant of salty ocean wind and is not easily af- on adaxial side scattered or in two lateral bands
fected by drought or pests. Its use as a hedge plant from base to apex; glands absent or inconspicuous,
has largely been superseded by the vigorous garden inactive; leaf colour dull green, rarely grey-green.
hybrid Leyland cypress, of which Monterey cypress Pollen cones numerous, terminal, solitary, ovoid-
is one of the parents. Several cultivar forms, notably oblong, 3–5 × 2 mm, more or less quadrangular or
with yellow-green foliage, have been developed. It is terete; micro­sporophylls (8–)10–14(–16), decus-
of interest to note that this is one of the two conifers sate, peltate, flattened or convex, bearing 3–4 abax-
in nature (virtually) confined to Monterey County ial, ­angular-globose, yellow pollen sacs. Seed cones
312 in California, that are among the most widely plant- often grouped but not densely clustered, terminal,
ed conifers in the world; the other species is Pinus growing in two seasons to mature, globose to sub-
radiata. globose, brown or grey-brown cones 15–30 mm
diam., which are persistent and variably serotinous.
Bract-scale complexes 6–8(–10), decussate, slowly
Cupressus sargentii Jeps., Fl. Calif. 1 (1): 61. 1909. parting, but the lowest pairs often remaining con-
Callitropsis sargentii (Jeps.) D. P. Little, Syst. Bot. nate; the ultimate pair in cones with 10 scales usually
31 (3): 474. 2006, (nom. ut. rej., Art. 56); Hespero­ very reduced; other scales peltate, 4–5 sided, angular,
cyparis sargentii (Jeps.) Bartel, Phytologia 91 (1): abaxially convex with a more or less prominent boss
183. 2009. Type: USA: California, Mendocino Co., or without a boss except for a minute curved bract
Mayacamas Range, Red Mountain, W. L. Jepson tip; abaxial surface nearly smooth to strongly ru-
3027 (holotype JEPS). gose, when still growing with a large, upcurved boss,
usually disappearing in mature cones but sometimes
retained; adaxially abruptly constricted, grooved,
Etymology
brown, usually glaucous, with lighter coloured seed
This species was named after Charles S. Sargent scars. Seeds 80–120 per cone, 3–5 mm long and wide,
(1841–1927), Director of the Arnold Arboretum. irregular, angular, slightly flattened, brown, lustrous,
often glaucous, with a lighter hilum; wings 2, form-
ing a thin strip 0.5(–1) mm wide around the seed.
Vernacular names
Sargent Cypress
Distribution
USA: California, Coast Ranges, divided into a
Description
northern and a southern group of populations. The
Trees to 20–25(–30) m tall, usually not taller than northern group is more widespread and is mostly
ca. 10 m; monopodial; trunk straight, or short, con- found north of the San Francisco Bay area to ca, 40°
torted, branching low, up to 100–120 cm diam. Bark N; the southern group occurs mostly in the Santa
on trunks to 4 cm thick, deeply fissured, with ana­ Lucia Range. The distance between the two main
stomosing ridges, fibrous, exfoliating in elongated groups is more than 250 km.
strips, grey-brown to grey. Branches spreading near- TDWG codes: 76 CAL
ly horizontally, lower branches often down-curved,
persisting along much of the trunk, foliage dense
Ecology
and bushy, forming a pyramidal or more rounded
or irregular crown. Foliage branches spreading or In the Upper Sonoran Life Zone, occasionally up to
ascending, ultimate branchlets irregularly disposed, the Transition Life Zone, in chaparral or pine-oak
rigid, 3–10(–15) × 1.2–1.6 mm, quadrangular or toru- woodland associated with Pinus attenuata, P. sabini-
lose, covered with scale leaves. Leaves decussate, ana, P. lambertiana, Pseudotsuga menziesii, Quercus
imbricate, appressed, on ultimate branchlets broadly spp., Arbutus menziesii, Arctostaphylos spp., Cea­
rhombic to more or less triangular, 1–2 × 1–1.2 mm, nothus spp., Adenostoma sp., and Yucca whipplei; on
gibbous or keeled below the obtuse or acute apex, dry hillsides, in ravines and canyons in clay, loam or
on whip shoots to 10 × 3 mm, with free, acute-­ sandy soil over serpentine. The altitudinal range is
from 50 m to 915 m a.s.l. The climate is of the Medi- decussate, imbricate, decurrent, (nearly) mono­
terranean type with hot, dry summers and winter morphic, scale-like, on ultimate branchlets 1–1.8
rain. mm long and rhombic, slightly gibbous, with ap-
pressed or free obtuse or acute; margins minutely
denticulate-hyaline; glands on all scale leaves; sto-
Conservation
mata few, scattered primarily on margins near leaf
IUCN: LC bases; leaf colour greyish green or green, sometimes
covered in thick cuticular wax. Pollen cones solitary
and terminal on ultimate branchlets, ovoid to ovoid-
Uses
oblong, 3–6 × 2–3 mm; microsporophylls 8–16,
No uses have been recorded of this species; in the decussate, bearing 3–6 abaxial pollen sacs near the 313
past farmers and ranchers may have utilised the lower margin. Seed cones solitary or grouped on lat-
wood for fenceposts. eral branches, terminal, maturing in 2 growing sea-
sons, persistent, subglobose to ovoid-oblong when
closed, (15–)20–35(–40) × (15)20–25(–30) mm, ma-
Cupressus sempervirens L., Sp. Pl. 2: 1002. 1753. turing to light brown or reddish brown with mostly
Type: Greece: Crete, [“Habitat in Creta”], Herb. parting scales. Bract-scale complexes (8–)10–14 in
Clifford 449 (lectotype BM). decussate pairs, of more or less equal size except for
smaller proximal (connate) and distal pairs, peltate,
Cupressus horizontalis Mill., Gard. Dict., ed. 8: Cu- poly­gonal (5–6-angular) in outline; abaxial surface
pressus No. 2. 1768; Cupressus sempervirens L. var. flat, convex or bossed, with protruding small bract
horizontalis (Mill.) Loudon, Hort. Brit. 1: 388. 1830; tip, (finely) rugose, abruptly narrowing towards
Cupressus sempervirens L. subsp. horizontalis (Mill.) cone axis; adaxial surfaces grooved, dark brown
A. Camus, [Les Cyprès] Encycl. Econ. Sylvicult. 2: with whitish seed marks near the base. Seeds 8–20
33. 1914. on each scale, closely packed, flattened, 3–6 × 3–4
mm, brown or red-brown with whitish hilum at
base; wings 2 on opposite sides, more or less equal
Etymology
in size and shape, as narrow strips surrounding the
The Latin species epithet means ‘evergreen’; Lin- seed, 0.6–1 mm wide.
naeus also classified Taxodium distichum, which is
deciduous, as belonging to his genus Cupressus.
Taxonomic notes
Traditionally, the Mediterranean Cypress has been
Vernacular names
divided into two elements, one with a fastigiate and
Mediterranean Cypress (and numerous local names one with a horizontally growing habit. The name
in different languages) Cupressus horizontalis Mill. has been given to trees
with a (horizontally) spreading branching habit:
Miller clearly distinguished the two growth forms
Description
as species. Various authors, as recently as Grilli
Trees to 40 m tall (usually smaller); monopodial; Caiola et al. (2000) have recognized the ‘horizontal
trunk to 2 m d.b.h. Bark eventually becoming thick, form’ at an infraspecific rank under C. sempervirens.
deeply fissured, hard, grey-brown, exfoliating slowly These authors accept C. sempervirens var. pyrami-
in small strips. Branches long, spreading or ascend- dalis for the fastigiate form, but if it is meant that
ing, sometimes fastigiate (the familiar ‘Italian cy- this includes the type of C. sempervirens that name
press’ of the Mediterranean is a selected/cultivated is illegitimate. Cupressus pyramidalis Targ.-Tozz. in
fastigiate form), forming a conical or pyramidal or Ann. Mus. Imp. Fis. Firenze 2 (2): 73 (1810) pertains
in old trees more or less sympodial, irregular and to the widely cultivated fastigiate growth form com-
broad, dense crown. Foliage branches spreading mon throughout the Mediterranean. That form is
or drooping, slender, ultimate branchlets subterete considered to be a cultivated form (‘cultigen’) and
to slightly flattened in cross section, 1–1.5(–2) mm not a taxon.
diam., slightly torose, persistent. Leaves covering
 intervals’ (W. T. Stearn, Botanical Latin ed. 3, 1983),
Distribution
and is perhaps referring to the ultimate branchlets.
E Mediterranean: Crete, Cyprus, East Aegean Is.,
Greece (?); N Africa: Libya; Western Asia: Iran, Is-
Vernacular names
rael, Jordan, Lebanon, Syria, and Turkey [W Medi-
terranean distribution based on cultigens]. Himalayan Cypress; Gulla, Gubraî (India, Nepal); xi
TDWG codes: 13 KRI 20 LBY 34 CYP EAI IRN LBS- zang bai mu (Chinese)
LB LBS-SY PAL-IS PAL-JO TUR
Description
Ecology
314 Trees to 35–40 m tall; trunk monopodial, up to 1.5 m
In maquis and in pine or juniper woodland associ- d.b.h. Bark smooth, orange-brown to red-brown,
ated with Pinus brutia, Juniperus excelsa, J. foetidis- becoming fibrous, greyish brown, exfoliating in
sima, J. drupacea, J. phoenicea, Quercus spp., Pistacia long strips. Branches spreading, assurgent or as-
atlantica, Amygdalus scoparia, and Poterium spino- cending, often ending drooping, forming a conical
sum; in rocky soil mostly over limestone on slopes or pyramidal or in old trees irregular and broad,
and in gorges, occasionally igneous rock. In its natu- dense crown. Foliage branchlets alternating, irregu-
ral habitat is occures from 90 m to 1700 m a.s.l. The larly disposed, spreading, lax or rigid depending on
climate is Mediterranean with dry, hot summers and thickness, quadrangular to subterete in cross sec-
winter rain, or semi-arid in more interior (eastern) tion, 0.8–1.5 mm diam., torose, persistent. Leaves
parts of its range. covering branchlets monomorphic, scale-like, on
ultimate branchlets 1–1.8 × 0.7–1.3 mm and rhom-
Conservation bic-gibbous, on whip shoots up to 5 mm long, with
incurved and appressed obtuse apex; margins entire
IUCN: LC
or minutely erose-hyaline; glands inconspicuous or
absent; stomata few, scattered primarily on margins
Uses
near leaf bases; leaf colour green or glaucous green.
There is a long history of exploitation going back to Pollen cones solitary and terminal, oblong, 3–6(–8)
the times of the ancient East Mediterranean and Le- × 1.5–2.5 mm; microsporophylls (8–)12–18(–20),
vantian civilisations. First, its wood was valued for decussate, bearing 3–4(–5) abaxial pollen sacs. Seed
its resistance to decay, later it also became an orna- cones solitary or in groups near upper ends of lateral
mental. This latter use has led to widespread intro- branches, terminal, maturing in 2 growing seasons,
duction throughout the Mediterranean at least from persistent, (sub)globose when closed, 12–20(–22)
Roman times to the present. In many villages and × 10–18 mm, maturing to (purplish) brown with
towns the fastigiate, columnar, or conical form is a parting scales. Bract-scale complexes (8–)10–12 in
very characteristic feature of the coastal and urban decussate pairs, peltate, polygonal (4–5-angular) in
landscapes. In several localities it regenerates spon- outline, nearly flat or more often bossed, with pro-
taneously, but the fastigiate habit betrays its cultivat- truding small bract tip, rugose, abruptly narrowing
ed origin. These old cultivars can be quite hardy and towards cone axis, with whitish seed marks near
withstand snow and frost to –20 °C or even lower. base. Seeds (4–)6–8 on each scale, ovoid-triangular,
A few modern cultivars, some dating from the 19th slightly flattened, 3–5 × 2–3 mm, brown or yellowish;
century, are known but rare in cultivation. wings 2 on opposite sides, more or less equal in size
and shape, to 1.2 mm wide.

Cupressus torulosa D. Don, in Lambert, Descr.

Taxonomic notes
Pinus 2: 18. 1824.
In recent years, reports of a cypress occurring in
Lang Son Province in the far northeast of Vietnam
Etymology
have been made from botanical surveys. Sometimes
The species epithet (from Latin torosus or torulosus) the name Cupressus tonkinensis Silba is applied to
means ‘cylindrical with bulges or contractions at these. Trees found there appeared to be planted,
or did occur in secondary vegetation giving rise to religious wood carving. Accessible stands are rare
suspicion about the indigenity of this plant. In Flora and in these straight trees have often been removed
du Cambodge, Laos et Vietnam 28: 79. (1996) this long ago. This species has been introduced in various
taxon is treated as C. torulosa D. Don “originaire de countries in Asia, either as an amenity tree in temple
l’Himalaya occidental”, but occuring on limestone grounds and monasteries or later as a plantation for-
hills between 250–1500 m a.s.l. in N Vietnam (and estry tree. It is also used in horticulture, especially
in S China?), presumably naturalized from intro- in southern Europe, where it is well established in
duction in the past but not explicitly designated as larger gardens and parks. A limited number of culti-
such. Native people appear to have cut most trees vars is known of this species.
due to a high demand for ‘incense’ from across the
Chinese border, so that perhaps only a few shrubs 2 varieties are recognized: 315
clinging to steep rocks presently survive. As I had
concluded earlier, they are morphologically identi- Cupressus torulosa D. Don var. torulosa. Cupressus
cal with C. torulosa var. torulosa (Farjon, 2005a) and lusitanica Mill. subsp. torulosa (D. Don) Silba,
most likely represent an introduced and naturalized Acta Phytol. Yunnanica 28 (5): 470. 2006. Type:
occurrence of that taxon far to the east of its natural India: Himalaya, [“Sooreh”], W. S. Webb W 6046A
range. (lectotype K-W). Fig. 99

Cupressus austrotibetica Silba, Phytologia 65: 334.

Distribution
1988.
Himalaya, from the Indus to the Brahmaputra; Chi- Cupressus karnaliensis Silba, J. Int. Conifer Preserv.
na: SE Xizang [Tibet], Yunnan. Soc. 1 (1): 19. 1994.
TDWG codes: 36 CHC-YN CHT 40 EHM-AP EHM- Cupressus karnaliensis Silba var. mustangensis Silba,
BH EHM-DJ EHM-SI IND NEP WHM-JK 41 VIE J. Int. Conifer Preserv. Soc. 1 (1): 22. 1994.
Cupressus tonkinensis Silba, J. Int. Conifer Preserv.
Soc. 1 (1): 23. 1994.
Ecology
Cupressus tongmaiensis Silba, J. Int. Conifer Preserv.
In the Himalaya, Cupressus torulosa is a codominant Soc. 1 (1): 24. 1994.
with Juniperus in the dry inner valleys and semi-arid Cupressus tongmaiensis Silba var. ludlowii Silba,
high mountain environments towards the Tibetan J. Int. Conifer Preserv. Soc. 1 (1): 24. 1994.
side of the main range, where this open forest or Cupressus pakistanensis Silba, J. Int. Conifer Preserv.
woodland type occupies S-facing slopes. The altitu- Soc. 15 (1): 4. 2008.
dinal range is 1560–3670 m a.s.l. Annual precipita-
tion does not exceed 300 mm, most of which falls
Description
in summer and autumn. North of the Himalaya,
in the Tibetan valleys that drain south through the Trees to 35 m tall, up to 1.5 m d.b.h. Ultimate branch-
Himalaya chain or north and east into the Zhangbo lets lax or rigid depending on thickness, 0.8–1.5 mm
system, conditions are too dry, but eastwards along diam., torose, persistent. Leaves on ultimate branch-
the Zhangbo River downstream from Gyangze lets 1–1.8 × 0.7–1.3 mm, rhombic-gibbous, glandular
C. torulosa is again found in isolated stands. Here we or eglandular; glands inconspicuous, in central de-
also find C. torulosa var. gigantea which is perhaps pression, elliptic.
a xeromorphic form of C. torulosa (S. Miehe, pers.
comm. May 2001, who claims to have seen both
Taxonomic notes
forms growing together).
For comments on several of Silba’s species names,
here cited in synonymy, see Farjon (2005a).
Uses
As of all Cupressaceae in Asia, the wood of this spe-
Distribution
cies is valued for many uses, primarily to do with its
durability (rot resistance). Traditionally it has been Himalaya: from the Indus (or N Pakistan?) to the
used for the construction of Buddhist temples and Brahmaputra; China: SE Xizang [Tibet].
 TDWG codes: 36 CHC-YN CHT 40 EHM-AP EHM-
Distribution
BH EHM-DJ EHM-SI IND NEP WHM-JK
China: S Xizang [Tibet] (Zangbo River Valley, from
ca. 93° to ca. 96° E); extreme NW Yunnan (vicinity
Conservation
of Dêqên on the Langcang [Mekong] River).
IUCN: NT TDWG codes: 36 CHC-YN CHT

Conservation
Cupressus torulosa D. Don var. gigantea
(W. C. Cheng & L. K. Fu) Farjon, Monogr. Cupressa­ Due to the scarcity of timber in the region where
316 ceae & Sciadopitys: 224. 2005. Cupressus gigantea scattered groves of var. gigantea occur, there is con-
W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): tinuous pressure for exploitation. Regeneration is
85. 1975. Type: China: Xizang [Tibet], [“22 km often poor due to grazing of livestock. Several groves
from Jia-mei-xi, Lang-bei Dong”], Qinghai-Xizang are protected as ‘sacred forest’ by Buddhist monks
Exped. 3318 (holotype PE). and some of the largest trees are on grounds used as
a cemetery, where any cutting is prohibited.
IUCN: VU (A1d)
Description
Trees to 30–40 m tall, up to 3.5 m d.b.h. Ultimate
Uses
branchlets short, usually thick, quadrangular to sub-
terete in cross section, (1–)1.5–2 mm diam., torose, Timber and firewood are locally taken. Large trees
persistent. Leaves on ultimate branchlets 1–1.8 × are often venerated in local traditions and religion.
0.7–1.3 mm, rhombic-gibbous, glandular or eglan-
dular; glands inconspicuous, in central depression,
elliptic.
Dacrycarpus (J. J. Bennett) de Laub., J. Arnold Arbor. 50: 315. 1969. Podocarpus sect.
Dacrydioideae J. J. Bennett in J. J. Bennett & R. Brown, Pl. Jav. Rar. 41. 1838. Type:
Dacrycarpus dacrydioides (A. Rich.) de Laub. [Podocarpus dacrydioides A. Rich.]
(Podocarpaceae).

Bracteocarpus A. V. Bobrov & Melikyan, Byull.

Distribution
Moskovsk. Obshch. Isp. Prir., Otd. Biol. 103 (1):
58. 1998. Type: Bracteocarpus imbricatus (Blume) Continental SE Asia: Myanmar [Burma]; S & SW
A. V. Bobrov & Melikyan [Podocarpus imbricatus China; Indochina. Malesia: from Peninsular Malay- 317
Blume]. sia to New Ireland (PNG). SW Pacific: New Caledo-
nia; Vanuatu; Fiji; New Zealand.
Greek: dakryon = a tear; karpos = fruit.
Key to the species of Dacrycarpus
Description
Transitional forms between the two kinds of leaves
Dioecious or rarely monoecious evergreen shrubs described in the key may exist in a single tree; it is
or trees. Resin canals in leaves and seed cones. Bark therefore necessary to consider the ‘extremes’ of
hard, occasionally with lenticels, becoming scaly. both forms where available in young trees and ma-
Leaves trimorphic, with small scale leaves, acicular ture trees alike. The two kinds are: scale-like, sub­
leaves and flattened, linear-falcate leaves, spirally ulate or acicular (not wider than thick), and linear
inserted, decurrent at base, appressed to widely to S-curved, foliar (bilaterally flattened).
spreading; in some species distichous, flattened lin-
ear-falcate leaves on juvenile plants and on determi- 1a. All foliage branchlets with leaves of a similar
nate vegetative lateral shoots of mature plants, but kind, shape acicular or subulate, sometimes
these absent in other species. Small scale leaves on slightly S-curved, 1.6–6(–10) mm long, 0.4–1
fertile shoots and on determinate vegetative lateral mm wide 2
shoots in some species; acicular leaves in the same 1b. Leaves of different kind on different foliage
positions in other species, both leaf types not disti- branchlets at least in young trees: scale-like,
chous, variously appressed to spreading. Stomata on subulate to acicular and linear to S-curved, bi-
both sides of the leaves (leaves amphistomatic). Pol- laterally flattened and longer in young trees and
len cones single or in pairs on axillary short shoots, with some species also both kinds in mature
at first nearly globular but elongating to short cylin- trees 4
drical, to ca. 10 × 3 mm; microsporophylls spirally 2a. Involucral leaves at base of seed cone 6–11 mm
attached to a slender rachis, with triangular heads long, incurved, enclosing the receptacle and
and two basal pollen sacs containing bisaccate pol- part of the seed(s) D. cinctus
len. Seed cones single on axillary short shoots with 2b. Involucral leaves at base of seed cone 4–7 mm
scale leaves and often surrounded by an ‘involu- long, more or less straight, enclosing the recep-
crum’ of acicular leaves at ends of foliar branchlets, tacle only 3
consisting of several spirally arranged bracts; only 3a. Leaves 1.6–3 mm long, acicular. Receptacles 4–5
a single subterminal one becoming fertillized and mm long, colouring dark purple; seeds 6–8 mm
forming an inverted seed; the remainder becom- long D. compactus
ing fused and partly swollen, forming a verrucose 3b. Leaves 2–6 mm long, acicular to slightly S-
receptacle ripening to red or purple and becoming curved. Receptacles 3–4 mm long, not chang-
succulent. Seeds completely surrounded by the soft ing colour or perhaps becoming yellowish;
epimatium being partly fused with the fertile bract, seeds 5–6(–7) mm long D. expansus
forming a grooved or shallow crest on one side of 4a. Pollen cones 20–30 mm long; microsporophylls
apex. subulate. Involucral leaves at base of seed cone
6–13 mm long, incurved, enclosing the recepta-
9 species. cle and part of the seed(s) D. cumingii
 4b. Pollen cones 7–12 mm long; microsporophylls
Etymology
apiculate. Involucral leaves at base of seed cone
1–8 mm long, spreading or incurved, at most The species epithet (Latin cinctus = enclosed or en-
only enclosing the receptacle 5 circled) refers to the seed cones enclosed by involu-
5a. Pollen cones 7–10 mm long, 1 mm wide. In- cral leaves.
volucral leaves at base of seed cone 1–2 mm
long, leaving the 2–3 mm long receptacle free
Vernacular names
D. vieillardii
5b. Pollen cones 8–12 mm long, 2–3 mm wide. In- sareh (Sulawesi); djariha (and many other names,
volucral leaves at base of seed cone 2–8 mm New Guinea)
318 long, at least in part enclosing the 3–7 mm long
receptacle 6
Description
6a. Involucral leaves at base of seed cone 5–8 mm
long, entirely enclosing the blue to purple re- Dioecious shrubs or trees to 30 m tall; trunk up to
ceptacle D. kinabaluensis 1 m d.b.h., erect; branches contorted, spreading,
6b. Involucral leaves at base of seed cone 2–5 mm crown eventually more or less flattened. Bark exfo-
long, partly to nearly entirely enclosing the or- liating in small plates or strips, brown weathering
ange to red receptacle 7 grey or blackish. All foliage branches spreading to
7a. Involucral leaves at base of seed cone 2–3 mm erect and with leaves of similar kind, acicular and
long, spreading, leaving most of the recepta- more or less bifacially flattened. Leaves on all shoots
cle free; seeds 3.5–4 mm long, smooth with a spirally arranged, not or rarely somewhat distichous
notched apex. Largest (foliar) leaves 3–7 mm on terminal vegetative shoots in young plants, all
long D. dacrydioides spreading slightly and equally around the shoot,
7b. Involucral leaves at base of seed cone 3–5 mm acicular (thin and hair-like on seedlings), curved
long, incurved, enclosing part or nearly all of inward, keeled abaxially, 2–6(–10) × 0.4–0.8 mm,
the receptacle; seeds 4–7 mm long, grooved with curved, apiculate apex. Leaves of branchlets
towards a curved apex. Largest (foliar) leaves in crowns of mature trees shorter (2–4 mm) than
(3–)5–12(–17) mm long 8 on younger trees, spreading at ca. 45°, decurrent,
8a. Foliar, more or less S-curved, distichously ar- curved inward, apiculate, often glaucous. Stomata
ranged leaves present in saplings and young on all sides (leaves amphistomatic) but on abaxial
trees, 5–10 mm long, 0.8–1.1 mm wide. Recep- side restricted to basal part of leaf, in 2 or more rows
tacles 3–4 mm long D. steupii on adaxial face up to near apex. Pollen cones termi-
8b. Foliar, more or less S-curved, distichously ar- nal on short or long shoots, subtended by acicular
ranged leaves present both in young trees and in leaves, nearly globose when immature, at maturity
mature trees, (3–)7–12(–17) mm long, 1–2 mm elongating to 8–10 mm long and 2–3 mm wide.
wide. Receptacles 4–7 mm long D. imbricatus Microsporophylls with apiculate apex, ca. 1.5 × 0.6
mm, each bearing two protruding pollen sacs. Seed
cones terminal on short shoots with spreading, 2–3
Dacrycarpus cinctus (Pilg.) de Laub., J. Arnold mm long, curved acicular leaves; involucral leaves
Arbor. 50: 332. 1969. Podocarpus cinctus Pilg., Bot. conspicuously longer, 6–11 mm, enclosing recepta-
Jahrb. Syst. 69: 253. 1938; Bracteocarpus cinctus cle and part of seed. Ripe receptacle 3–4 mm long,
(Pilg.) A. V. Bobrov & Melikyan, Bjull. Moskovsk. warty, red or purplish tinged. Ripe seeds 1(–2) on a
Obsc. Isp. Prir., Otd. Biol. 103 (1): 58. 1998. Type: receptacle, subglobose, 4–6 mm long, including the
Papua New Guinea: Morobe, Búsú River, Mt. smooth, light or dark red-brown, sometimes glau-
Sarawaket, M. S. Clemens 5261 (holotype B, cous epimatium, with a longitudinal grooved crest
destroyed; isotypes A, LAE, NY). Fig. 100 terminating in a curved protruding apex.

Podocarpus dacrydiifolius Wasscher, Blumea 4 (3):

Distribution
410. 1941; Bracteocarpus dacrydiifolius (Wasscher)
A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc. Malesia: Borneo (Sarawak), Sulawesi, Maluku
Isp. Prir., Otd. Biol. 103 (1): 59. 1998. [Moluccas] (Ceram); Papuasia: New Guinea.
TDWG codes: 42 BOR-SR MOL SUL 43 NWG-IJ
Vernacular names
NWG-PN
kaipik, umbwa and many other names varying with
the languages of native tribes
Ecology
Dacrycarpus cinctus occurs from montane rain­forest
Description
at around 1800 m to alpine low mossy forest or shrub-
land and tree fern grassland in New Guinea as high Dioecious trees to 20 m tall; trunk up to 60 cm
as 3600 m a.s.l. In Sulawesi it is reported from lower d.b.h., erect or contorted; branches irregular and
montane forest at 900 m. In high montane forest it contorted; crown eventually more or less flattened.
attains tall tree size and is dominant (emergent) or Bark exfoliating in small plates or strips, dark brown 319
codominant with Nothofagus, Elaeocarpus, and the weathering grey; inner bark reddish. All foliage
conifers Papuacedrus papuana and Podocarpus spp. branches spreading to erect; terminal vegetative
(in New Guinea). Above 3000 m Cunoniaceae and shoots not distichous in young plants. Leaves on
Myrtaceae become abundant and the forest patches all shoots of similar kind, acicular and more or less
are often interspersed with swampy and peaty tus- bifacially flattened, spirally arranged, imbricate and
sock grasslands dominated by solitary clumps of spreading equally on all sides of the shoot, acicular
Dacrycarpus and scattered tree ferns (Cyathea) un- (thin and hair-like on seedlings), curved inward,
til the last shrubby specimens reach the tree line as strongly keeled abaxially, 1.6–3 × 0.6–1 mm, with
solitary shrubs or krummholz trees. acute apex. Leaves of branchlets in crowns of mature
trees virtually similar to those of young trees, slight-
ly wider at base, imbricate, short decurrent, curved
Conservation
inward, acute to apiculate, often glaucous. Stomata
IUCN: LC on all sides (leaves amphistomatic) but on the abax-
ial side restricted to basal part of leaf, in 2 or more
rows on the adaxial face up to near the apex. Pollen
Uses
cones terminal on short or long shoots, subtended
Large trees of this species will be valuable timber by acicular leaves, nearly globose when immature, at
and have undoubtedly been logged for this purpose. maturity elongating to 8–10 mm long and 2–3 mm
Its wood is probably not distinguished from other wide. Microsporophylls with apiculate apex, 1.5–2
members of the family and traded as ‘podocarp’ × 0.5 mm, yellow with reddish apex, each bearing
wood. Its properties and uses would be similar to two protruding pollen sacs. Seed cones terminal on
those of D. imbricatus. In Sarawak and Sulawesi the short shoots with imbricate, 1.6–3 mm long, curved
wood has been traditionally used for the construc- acicular leaves; involucral leaves longer, 5–7 mm, en-
tion of longhouses. closing the receptacle and reaching but not enclos-
ing the seed. Ripe receptacle 4–5 mm long, warty,
green turning dark purple. Ripe seeds 1(–2) on a
Dacrycarpus compactus (Wassch.) de Laub., J. Ar- receptacle, subglobose, 6–8 mm long, including the
nold Arbor. 50: 336. 1969. Podocarpus compactus smooth, green to purplish brown epimatium, with a
Wasscher, Blumea 4 (3): 411. 1941; Bracteocarpus longitudinal grooved crest terminating in a curved
compactus (Wasscher) A. V. Bobrov & Melikyan, protruding apex.
Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1):
58. 1998. Type: Papua New Guinea: Owen Stanley
Taxonomic notes
Range, Mt. Albert Edward, L. J. Brass 4284 (holo-
type L). This species has formerly been known as Podo­
carpus papuanus, a name given to another species
by Ridley in 1916 and now a synonym of Dacry­
Etymology
carpus imbricatus var. robustus. Wasscher named the
The species epithet refers to the dense and compact present species Podocarpus compacta (which should
foliage of older trees. have been the masculine form compactus) and it
was subsequently placed in the genus Dacrycarpus
 by David de Laubenfels in 1969, where subsequent Dacrycarpus cumingii (Parl.) de Laub., J. Arnold
accounts have all accepted it to belong. The genus Arbor. 50: 329. 1969. Podocarpus cumingii Parl., in
name Bracteocarpus when proposed by Bobrov & Candolle, Prodr. 16 (2): 521. 1868; Bracteocarpus
Melikyan included the type of Dacrycarpus de Laub. cumingii (Parl.) A. V. Bobrov & Melikyan, Bjull.
and is therefore an illegitimate name. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 58.
1998. Type: Philippines: Luzon, Tayabas Prov.,
H. Cuming 803 (syntype K).
Distribution
New Guinea (highlands).
Etymology
TDWG codes: 43 NWG-IJ NWG-PN
320 This species was named in honour of the plant col-
lector H. Cuming, who collected the specimens on
Ecology
which the original description was based.
Dacrycarpus compactus is a highland species occur-
ring in subalpine shrubberies and on the fringes of
Vernacular names
alpine tussock grassland dominated by Deschamp-
sia klossii. It is common in coniferous high montane sangu (Sumatera); igem (Phillipines: Davao, Mind-
forest with Papuacedrus papuana, Podocarpus spp., anao) and other local names.
and a few angiosperms and then becomes more
abundant and often a dominant emergent tree in
Description
mossy low forest and shrubbery fringing peaty wet
tussock grasslands. Here the deepest peat is grass- Dioecious trees to 38 m tall; trunk up to 1 m d.b.h.,
land and the conifers Dacrycarpus and Papuacedrus erect, monopodial; crown eventually spreading,
occupy stony rises and scree slopes where the peat is dome-shaped. Bark exfoliating in small plates or
thinner or absent and drainage better. Isolated trees strips, brown weathering grey. Foliage branches
may occur scattered in the grassland as they are rela- with leaves of two kinds, more or less acicular and
tively resistant to grass fires once they reached some flattened. Primary shoots of seedlings slender, of
size. There is a cool, misty climate at these heights mature trees slender or in exposed crowns short and
that has almost no seasonal changes. The altitudinal stiff, terminating in a bud-like cluster of incurved
range of this species is (2800)3000–4300 m a.s.l. acicular leaves. Leaves on all shoots spirally ar-
ranged, subulate or acicular (thin and hair-like on
seedlings), curved inward at tip, spreading slightly,
Conservation
keeled abaxially, with curved, apiculate apex. Leaves
IUCN: LC on 2–6 cm long terminal and deciduous branchlets
of seedlings to young trees flattened, distichous,
decurrent, linear or slightly s-curved, with parallel
Uses
smooth margins, 5–14 mm long (shortest at base and
This tree of moderate size produces useful timber of apex of branchlet), 0.8–1.3 mm wide, weakly keeled
fine quality but its inaccessibility and relative scar- on both surfaces; apex curved forward, apiculate.
city of straight trees of good size prevent commer- Leaves of branchlets in crowns of mature trees simi-
cial exploitation. The species is not in cultivation for lar to more acicular, 2–4(–6) × 0.4–0.7 mm, less dis-
forestry or horticulture. tichously arranged to spreading on all sides, curved,
apiculate. Stomata on both kinds of leaves and on
all sides (leaves amphistomatic) in many intermit-
tent rows on flat, larger leaves, in 2 or more rows on
each face of acicular leaves. Pollen cones terminal
on short shoots, subtended by small acicular leaves,
short when immature, at maturity elongating to 20–
30 mm long and 2–3 mm wide. Microsporophylls
subulate like leaves, ca. 1.5 × 0.5 mm, each bearing
two protruding pollen sacs. Seed cones terminal on
short shoots with spreading, 7–13 mm long, curved Dacrycarpus dacrydioides (A. Rich.) de Laub.,
acicular leaves enclosing receptacle and seed. Ripe J. Arnold Arbor. 50: 337. 1969. Podocarpus dacry­
receptacle 3–4 mm long, warty, reddish. Ripe seeds dioides A. Rich., in Lesson & Richard, Voy. Astro-
1(–2) on a receptacle, subglobose, 3.5–4.5 mm long, labe Bot. [Ess. Fl. Nouv. Zél.] 1: 358. 1832. Type: New
including the smooth, light yellowish brown to Zealand: North Island, J. S. C. Dumont d’Urville s.n.,
blackish brown, sometimes glaucous epimatium, 1827 (holotype? G-DC). Fig. 101, 102, 103
with a longitudinal grooved crest terminating in a
curved protruding apex.
Etymology
The species epithet alludes to a similarity with
Distribution
Dacrydium. 321
Malesia: Borneo (Sarawak), Philippines, N Sumat-
era (Gunung Leuser N. P.).
Vernacular names
TDWG codes: 42 BOR-SR PHI SUM
White pine; kahikatea (Maori)
Ecology
Description
Dacrycarpus cumingii occurs in mossy forest. The
climate is cool and wet and fog is shrouding the Large dioecious trees to 60 m tall; trunk 1.5–2 m
mountains much of the time. Lowest records are d.b.h., erect, monopodial, old trees often butressed
from 1600 m a.s.l. in montane forest, where more near the base. Bark smooth at first, becoming rough
commonly D. imbricatus would occur and where and scaly, exfoliating with large flakes, brown turn-
both species may be associated with Agathis spp. ing grey; inner bark reddish. Crown conical in
and Sundacarpus amarus, the highest from 3300 m young trees, becoming domed or spreading in old
a.s.l. near the tree line. trees with heavy lower branches. Foliage branches
with leaves of three kinds, short acicular, inter­
mediate and longer flattened. Leaves on all branches
Conservation
of seedlings and saplings to 1–2 m tall bilaterally
IUCN: LC flattened, distichous, decurrent, slightly S-curved,
with parallel smooth margins, 3–7 mm long (short-
est on primary branches and at base and apex of
Uses
the terminal branchlets), 0.6–1 mm wide; apex
This species can grow to a tall forest tree and such curved forward, apiculate. Intermediate leaves ap-
specimens yield valuable timber, but it is usually pearing on young but often fertile trees on primary
smaller and grows at high altitude where timber and terminal branches, spirally arranged, not dis-
extraction is more difficult. The uses of its wood in tichous, spreading at ca. 45°, 2–4 mm long. Leaves
Borneo are assumed to be similar to that of D. im- on primary shoots, fertile shoots, and some termi-
bricatus where it is available in low quantities, e.g. nal shoots of large mature trees spirally arranged,
construction for local houses, furniture and tool mostly sub­ulate or scale-like, curved inward at the
making. In the Philippines where D. cumingii is (or tip, appressed to imbricate, keeled abaxially, 1–2 ×
was) more common it is an important source of face 0.4–1 mm, apiculate. Stomata on all kinds of leaves
veneer. and on all sides (leaves amphistomatic) in several
intermittent rows from base to apex. Pollen cones
terminal (or paired) on short scale-leaved shoots,
at first semi-globose but elongating to 8–10(–12) ×
2 mm; microsporophylls triangular, incurved, 1–1.2
× 0.8 mm, apiculate, green, with two light yellow
pollen sacs at their base. Seed cones terminal on
short shoots with spreading, 2–3 mm long sub­ulate
leaves enclosing the initial structure, subtending the
receptacle later. Receptacle 5–7 mm long, swelling
 to ­subglobose shape, warty, turning from (green) (I experienced from a private landowner first hand
yellow to orange to red, with only tips of 1–3 bract that he was “not allowed to touch these trees” but I
leaves protruding. Ripe seeds 1 on a receptacle, was free to photograph them) and consequently its
subglobose, 3.5–4 mm long, including the smooth, wood is no longer traded. In New Zealand it is not
lustrous, purple-­glaucous epimatium, with a barely cultivated as an ornamental tree (New Zealanders
visible notched apex. prefer to plant ‘British’ gardens not native ones) and
it would require a mild, moist climate to do so. Its
profuse and colourful ‘fruit’ (only present in female
Distribution
trees) and delicate foliage are certainly attractive, but
New Zealand. it is slow growing. It is in cultivation in SW England
322 TDWG codes: 51 NZN NZS and Ireland in some of the famous woodland gar-
dens and could certainly be tried elsewhere if given
proper growing conditions.
Ecology
This still widespread species used to be a dominant
emergent tree in lowland podocarp forests which Dacrycarpus expansus de Laub., J. Arnold
have largely been destroyed by unlimited logging Arbor. 50: 334. 1969. Podocarpus expansus (de
and forest clearance for agriculture. It is now virtu- Laub.) Whitmore, in Whitmore et al. (eds.),
ally restricted to forest reserves on steeper terrain at Tree Fl. Indonesia, Checkl. Irian Jaya: 234. 1997;
altitudes up to 600 m a.s.l. These are mixed coni- Bracteocarpus expansus (de Laub.) A. V. Bobrov
fer-angiosperm forests with usually dominance of & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
Agathis australis (only in the far north of New Zea- Otd. Biol. 103 (1): 59. 1998. Type: Papua New
land), and/or several of the ‘podocarps’ Dacrydium Guinea: Western Highlands, Laiagam Subdistrict,
cupressinum, Podocarpus totara, P. cunninghamii, [“Yobobos Grassland”], R. D. Hoogland &
Prumnopitys taxifolia, P. ferruginea, Halocarpus bid- R. Schodde 7463 (holotype L).
willii, Manoao colensoi, and Phyllocladus trichoma-
noides. Angiosperm trees in these forests are e.g.
Etymology
Beilschmiedia tarairi, Laurelia novae-zelandiae (in
swampy places), Metrosideros robusta (which begins The species epithet refers to the spreading leaves on
life as an epiphyte), and especially in South Island foliage branches of adult trees.
Nothofagus spp. Shrubs and tree ferns (Cyathea,
Dicksonia) as well as epiphytes of many kinds can be
Vernacular names
abundant in these moist evergreen forests in a sub-
tropical to warm temperate climate. pau or pa’u in the Enga language

Conservation Description
IUCN: LC Dioecious trees to 25(–30) m tall; trunk up to 60 cm
d.b.h., erect or contorted; branches irregular, con-
torted, crown eventually more or less flattened. Bark
Uses
exfoliating in coarse plates or strips, dark brown
Extensively logged with other species and hardly weathering grey; inner bark pink to reddish brown.
distinguished in the timber trade from other ‘podo- Foliage branches spreading or sometimes pendu-
carp’ wood, White pine (now preferably called by its lous, with leaves of slightly different kind, juvenile
Maori name kahikatea) was an extremely valuable longer and S-curved leaves gradually disappearing in
tree due to its potential size and the good qualities of crowns of older trees. Leaves on primary shoots and
podocarp wood. It was used for carpentry, flooring fertile shoots spirally arranged, on lateral branchlets
and paneling in houses, dry cooperage, tool handles, often more or less distichous or 4-ranked especial-
etc. Today this species enjoys near total protection ly in young plants, acicular (thin and hair-like on
seedlings), curved inward, strongly keeled abaxi- Dacrycarpus imbricatus (Blume) de Laub.,
ally, 2–6 × 0.5–1 mm, with apiculate apex. Leaves of J. Arnold Arbor. 50: 317. 1969.
branchlets in crowns of mature trees similar to those
of young trees, subulate-acicular, not distichous or
Etymology
S-curved, and 2–3 mm long, spreading at 30–45°,
short decurrent, curved inward, acute to apiculate, The species epithet refers to the imbricate (overlap-
often glaucous. Stomata on all sides (leaves amphis- ping) leaves on primary shoots.
tomatic) but on the abaxial side restricted to basal
part of leaf, in 2 or more rows on the adaxial face up
Vernacular names
to near apex. Pollen cones terminal on short or long
shoots, subtended by acicular leaves, nearly globose Numerous vernacular names are given under four 323
when immature, at maturity elongating to 8–10 mm varieties by De Laubenfels (1988) in Flora Malesi-
long and 2–3 mm wide. Microsporophylls with long ana, but this is clearly based on geography, not on
apiculate apex, 1.5–2 × 0.5 mm, yellow with reddish the recognition of these varieties by local people. No
apex, each bearing two protruding pollen sacs. Seed single vernacular name is in use for the species as a
cones terminal on short shoots with imbricate, 2–3 whole, although Pilger (1903) cites two: kimerah and
mm long, curved acicular leaves; involucral leaves kiputri. The latter names seem to apply to ‘podo­
slightly longer, 4–6 mm, enclosing receptacle only. carps’ of Jawa, not to a particular species.
Ripe receptacle 3–4 mm long, warty, green or olive
green (?). Ripe seeds 1(–2) on a receptacle, subglo-
Description
bose, 5–6(–7) mm long, including the smooth, green
to purplish brown epimatium, with a longitudinal Shrubby to large dioecious trees to 40–50 m tall, up
grooved crest terminating in a curved protruding to 2 m d.b.h.; trunk erect, terete; branches of large
apex. trees spreading and ascending, forming a broad,
open crown. Bark hard, with rough surface, on large
trees breaking into thick, slightly elongate plates or
Distribution
exfoliating in short strips, dark brown weathering
New Guinea (mostly Central Highlands of Papua greyish white, grey or blackish; inner bark pink to
New Guinea). reddish brown and slightly fibrous. Foliage branches
TDWG codes: 43 NWG-IJ NWG-PN with leaves of two kinds, short acicular and longer
flattened. Leaves on primary shoots, fertile shoots
and some terminal shoots of mature trees spirally
Ecology
arranged, subulate or acicular (thin and hair-like
Dacrycarpus expansus occurs in lower montane to on seedlings), curved inward at tip, appressed to
high montane forests, often on the margins of tus- imbricate or spreading, keeled abaxially, 1–3(–4) ×
sock grassland with tree ferns (Cyathea). It can be 0.4–1 mm, apiculate. Leaves on many 1–5(–7) cm
mixed with Papuacedrus papuana or occur in nearly long terminal and deciduous branchlets bilaterally
pure stands. The soil is often water-logged, peaty flattened, distichous, decurrent, slightly S-curved,
and acidic. with parallel smooth margins, (3–)7–12(–17) mm
long (shortest at base and apex of branchlet), 1–2
mm wide; apex curved forward, apiculate; this type
Conservation
of foliage branchlets often persist in crowns of ma-
IUCN: LC ture trees. Stomata on both kinds of leaves and on
all sides (leaves amphistomatic) in 3–6 intermittent
rows on flat, larger leaves, in 1–2 rows on each face
Uses
of acicular leaves. Pollen cones terminal on short
Little is known about the uses of this species; it is shoots, subtended by small acicular leaves, nearly
assumed to be logged on a small scale for local use globose when immature, at maturity elongating to
mainly, and if traded, would be called ‘podocarp’ 8–12 mm long and 2–3 mm wide. Microsporophylls
together with species of Podocarpus logged in the with triangular, apiculate apex, ca. 1.2 × 0.8 mm,
Central Highlands. each bearing two protruding pollen sacs. Seed cones
 terminal on short shoots with spreading, 3–5 mm
Uses
long acicular leaves enclosing the initial structure,
subtending the receptacle later. Ripe receptacle 4–7 This widespread species is one of the most valuable
mm long, warty, orange-red or red, with protruding timber trees in SE Asia. Its wood, known as ‘po-
green bract leaves. Ripe seeds 1(–2) on a receptacle, docarp’ or ‘melur’ (Indonesia) in the timber trade,
subglobose, 5–7 mm long, including the smooth, is there not differentiated from other genera like
glaucous green to red-brown epimatium, with a Nageia and Podocarpus. Dacrycarpus imbricatus
longitudinal grooved crest terminating in a curved produces excellent pulpwood due to its relatively
protruding apex. long fibres, but its higher grades are put to more spe-
cialized uses such as furniture (especially table tops),
324 cabinet work, and wood carving in Thailand, the
Distribution
Philippines as well as in Fiji, masts for sailing boats,
China: Hainan, Guangxi, NW Yunnan; Indochina; panelling, and veneer, light construction ­timbers,
Malesia; Papuasia; SW Pacific: Fiji, Vanuatu. drawing boards, tea chests, utensils, and much
TDWG codes: 36 CHC-YN CHH CHS-GX 41 CBD more (ETI/PROSEA, CD-ROM on Timber trees;
LAO MYA THA VIE 42 BOR-BR BOR-KA BOR-SB www.eti.uva.nl). It is used in tropical countries as an
BOR-SR JAW LSI-LS MLY-PM PHI SUL SUM 43 BIS ornamental tree in parks and gardens.
NWG-IJ NWG-PN 60 FIJ VAN
3 varieties are recognized:
Ecology
Dacrycarpus imbricatus (Blume) de Laub. var.
This widespread species is growing in primary imbricatus. Podocarpus imbricatus Blume, Enum.
and secondary lower montane to upper montane Pl. Javae 1: 89. 1827; Bracteocarpus imbricatus
rainforests, common or scattered, codominant or (Blume) A. V. Bobrov & Melikyan, Bjull. Moskovsk.
rarely without codominants, it is a canopy tree or Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998. Type:
an emergent. Dacrycarpus imbricatus is common Indonesia: Jawa, [“western Java”], C. L. Blume s.n.
on volcanic or ultramafic soils, and occasionally (lectotype L). Fig. 104
occurs on sandstone or limestone. It occurs most
widely in Lithocarpus-Castanopsis mixed forest, of- Podocarpus kawaii Hayata, Bull. Econ. Indochine 19:
ten on steeper slopes and on ridges. In W Jawa var. 439. 1917; Bracteocarpus kawaii (Hayata) A. V. Bo-
imbricatus occurs on Mt. Tjeremai only with Podo- brov & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
carpus neriifolius and Altingia noronhae between Otd. Biol. 103 (1): 59. 1998.
2400 and 2700 m a.s.l. In Lombok D. imbricatus has Dacrycarpus imbricatus (Blume) de Laub. var. patu-
been found as low as 200 m while in Sulawesi it as- lus de Laub., J. Arnold Arbor. 50: 320. 1969.
cends to ca. 3000 m and in New Guinea to 3720 m.
At elevations above ca. 1200 m, i.e. above the dipte-
Description
rocarp rainforests of Malesia, Lithocarpus (tropical
oaks), Castanopsis (chestnuts) and other conifers Leaves on primary shoots of mature trees slightly
(Agathis, Dacrydium, Phyllocladus, Podocarpus) are spreading, more or less free, or sometimes imbricate
the important associated trees; in New Guinea it and mostly appressed (shoots appear very slender),
occurs also with Nothofagus and Phyllocladus hypo­ spreading only here and there, 1–2(–3) × 0.4–0.6
phyllus. On ultramafic substrates Casuarinaceae mm.
(e.g. Ceutho­stoma, Gymnostoma), Myrtaceae (e.g.
­Eugenia, Leptospermum, Tristania, Xanthomyrtus)
Taxonomic notes
and the conifer genus Dacrydium are the most com-
mon associates of Dacrycarpus imbricatus. The up- According to De Laubenfels var. patulus is restrict-
per montane forests above ca. 1800 m are rich in ed to Jawa, the Lesser Sunda Islands, and Sulawesi
epiphytes, especially mosses are very abundant and [Flora Malesiana ser. 1, 10 (3): 377, f. 25 (1988)] but
hang from every branch and these forests are often collections at K and FHO that seem to belong to
shrouded in fog for days on end. this form with slender, inbricate primary leaves
have been gathered from Thailand to Vanuatu and
Fiji, i.e. through the entire range of the species. The Dacrycarpus steupii de Laub., Kalikasan 7 (2): 127.
distinction between this form and branchlets with 1978, non de Laub. 1969.
more spreading primary leaves is often gradual and
both may well occur in the same tree. Verification
Description
of specimens at KEP (Forest Research Institute of
Malaysia) and observations in the field in Septem- Leaves on primary shoots robust, 2–6 mm long,
ber-October 2008 confirmed this. Dacrycarpus 0.6–1 mm wide, often nearly all spreading around
imbricatus var. patulus does not appear to be truly the shoot, involucral leaves spreading.
distinct and is here treated as a synonym of Dacry-
carpus imbricatus var. imbricatus.
Distribution
325
Malesia: Borneo, Maluku [Moluccas], Philippines;
Distribution
Papuasia: New Guinea.
As for the species. TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
MOL PHI 43 NWG-IJ NWG-PN
Conservation
Conservation
IUCN: LC
IUCN: LC
Dacrycarpus imbricatus (Blume) de Laub. var.
curvulus (Miq.) de Laub., J. Arnold Arbor. 50: 326.
1969. Podocarpus cupressinus R. Br. ex G. Benn. Dacrycarpus kinabaluensis (Wassch.) de Laub.,
var. curvulus Miq., Pl. Jungh. 1: 4. 1851. Type not J. Arnold Arbor. 50: 330. 1969. Podocarpus im-
designated (coll. by F. W. Junghuhn in Jawa). bricatus Blume var. kinabaluensis Wasscher, Blu-
mea 4 (3): 400. 1941; Bracteocarpus kinabaluensis
(­Wasscher) A. V. Bobrov & Melikyan, Bjull. Mos­
Description
kovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998.
Shrubby trees to 8 m tall; foliage branchlets of ma- Fig. 105, 106
ture trees drooping to pendulous, with adult type
imbricate scale leaves 1.2–3 × 0.8–1 mm; involucral
Etymology
leaves spreading but apically incurved, not exceed-
ing the receptacle in length. The species epithet means ‘from Kinabalu’, the
mountain on which it is endemic.
Distribution
Vernacular names
Indonesia: Jawa, N Sumatera.
TDWG codes: 42 JAW SUM No common names are recorded for this species.

Conservation Description
IUCN: LC Dioecious shrubs or small trees to 13 m tall, with
up to 30 cm stem diam.; trunk of trees short, often
Dacrycarpus imbricatus (Blume) de Laub. var. gnarled; crown becoming dense. Bark exfoliating
robustus de Laub., J. Arnold Arbor. 50: 323. 1969. in small plates or strips, brown weathering grey.
Type: Papua New Guinea: Eastern Highlands, Mt. Foliage branches with leaves of two kinds only on
Wilhelm, L. J. Brass 30568 (holotype A). seedlings and saplings, or on trees with adventitious
shoots. Leaves on primary shoots spirally arranged,
Podocarpus papuanus Ridl., Trans. Linn. Soc. Lon- more or less imbricate, subulate or acicular, curved
don, Bot., ser. 2, 9: 158. 1916; Bracteocarpus papuanus inward or more or less straight, keeled abaxially, 3–6
(Ridl.) A. V. Bobrov & Melikyan, Bjull. Moskovsk. × 0.7–1 mm, with curved, apiculate apex. Leaves on
Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998. 2–4(–5) cm long terminal (deciduous) branchlets of
 seedlings and saplings (or adventitious and shaded orchids and pitcher plants (Nepenthes spp.) are com-
shoots) flattened, distichous or nearly 4-ranked, mon, and epiphytes are abundant on the shrubs and
decurrent, linear or slightly S-curved, with paral- low trees.
lel smooth margins, 5–10 mm long (shortest at base
and apex of branchlet), 0.8–1.3 mm wide, more or
Conservation
less smooth; apex curved forward, apiculate. Leaves
on terminal branchlets in crowns of mature trees This species is only known from the higher parts
spirally arranged or sometimes 4-ranked, never dis- of Mt. Kinabalu and consequently occupies only a
tichously arranged, shorter and narrower, curved, small area probably less than 100 km² in extent. On
apiculate. Stomata on both kinds of leaves and on the other hand, being in the centre of one of the bet-
326 all sides (leaves amphistomatic) in 2–5 inter­mittent ter protected National Parks in the whole of SE Asia
rows on flat, larger leaves, in 1 grooved row on each provides it with a degree of protection, and no de-
face of acicular leaves. Pollen cones terminal on cline is known at present. Increased tourism might
short shoots, subtended by small acicular leaves, pose indirect threats to this species, for instance
nearly globose when immature, at maturity elongat- by increasing the incidence of fires in episodes of
ing to 8–12 mm long and 2–3 mm wide. Microsporo­ draught associated with El Niño climatic cycles.
phylls with triangular, apiculate apex, ca. 1.2 × 0.8 IUCN: NT
mm, each bearing two protruding pollen sacs. Seed
cones terminal on short shoots with spreading, 5–8
Uses
mm long, incurved acicular leaves enclosing the re-
ceptacle only. Ripe receptacle 4–5 mm long, warty, This species occurs within a National Park at high
blue or purple. Ripe seeds 1(–2) on a receptacle, altitude and is not used economically. As far as
subglobose, 5–7 × 5–6 mm mm long, including the known it is not in cultivation except perhaps in a
smooth, light yellowish brown to brown, sometimes few botanic gardens and/or private collections.
glaucous epimatium, with a longitudinal grooved
crest terminating in a curved protruding apex.
Dacrycarpus steupii (Wasscher) de Laub.,
J. Arnold Arbor. 50: 328. 1969. Podocarpus steupii
Distribution
Wasscher, Blumea 4 (3): 405. 1941; Bracteocarpus
Borneo: Sabah (Mt. Kinabalu). steupii (Wasscher) A. V. Bobrov & Melikyan,
TDWG codes: 42 BOR-SB Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103
(1): 59. 1998. Type: Indonesia: Sulawesi, Enrekang,
Rantelemo, leg. ign. NIFS 22857 (holotype not
Ecology
located, isotype BO).
Dacrycarpus kinabaluensis is a shrubby tree restrict-
ed to the upper montane forest and subalpine dwarf
Etymology
forest on Mt. Kinabalu (4101 m). It occurs on this
mountain from ca. 2600 a.s.l. m up to the tree line This species was named after Dutch forester
at ca. 3500 m. Dacrycarpus imbricatus occurs to at F. K. M. Steup, who collected plants in Sulawesi in
least 2400 m and possibly higher on this mountain, the 1930’s.
but is a tree of taller forest. Dacrycarpus kinabaluen-
sis is growing predominantly on ultramafic rock but
Vernacular names
becomes one of the dominant shrubs above 3000
m on granite. It can form dense, nearly pure stands miejoop, nak, pau and other names are used for this
but is commonly associated with other conifers, e.g. species in New Guinea
Dacrydium gracile, D. gibbsiae, Phyllocladus hypo-
phyllus and Podocarpus brevifolius. Angiosperm
Description
trees are often scarce but heather-like tall shrubs and
dwarfed trees are common with Rhododendron spp. Dioecious trees to 36 m tall; trunk up to 1 m d.b.h.,
and Leptospermum spp. the most abundant. There is erect, monopodial; crown more or less conical.
often a thick moss layer on the forest floor, in which Bark exfoliating in small plates or strips, brown
­ eathering grey. Foliage branches with leaves of two
w altitudes it forms clumps, sometimes without other
kinds, acicular leaves and flattened leaves. Leaves on trees present, or mixed with Papuacedrus papuana
primary shoots and fertile shoots spirally arranged, in New Guinea. The altitudinal range is from 860 m
imbricate or spreading, subulate or acicular (thin to 3470 m a.s.l.
and hair-like on seedlings), curved inward at tip,
keeled abaxially, 2–3(–4) × 0.5–0.8 mm, with curved,
Conservation
apiculate apex. Leaves on 2–5 cm long terminal and
deciduous branchlets of seedlings to young trees bi- Dacrycarpus steupii has been collected from Gu-
laterally flattened, distichous, decurrent, linear or nung Beratus, a mountain near Balikpapan, but
slightly s-curved, with parallel smooth margins, 5–10 it had disappeared there by the early 1980’s due to
mm long (shortest at base and apex of branchlet), deforestation. It is still present in central Sulawesi 327
0.8–1.1 mm wide, weakly keeled on both surfaces; and the Philippines and widespread in New Guinea;
apex curved forward, apiculate. Leaves of terminal consequently the global status of this species is still
branchlets in crowns of mature trees similar to more considered outside the threatened categories.
acicular, shorter (2–6 mm) and narrower, 4-ranked IUCN: NT
to more commonly spirally arranged, decurrent,
curved inward, apiculate. Stomata on both kinds of
Uses
leaves and on all sides (leaves amphistomatic) in 4–6
intermittent rows in two bands on flat, larger leaves, Large trees will be valuable timber and have un-
in 1–2 rows on each face of acicular leaves. Pollen doubtedly been logged for this purpose. Its wood is
cones terminal on short shoots, subtended by small probably not distinguished from other members of
acicular leaves, nearly globose when immature, at the family and traded as ‘podocarp’ wood. Its prop-
maturity elongating to 8–12 mm long and 2–3 mm erties and uses would be similar to those of D. im-
wide. Microsporophylls with triangular, apiculate bricatus and particularly useful for construction of
apex, ca. 1.2 × 0.8 mm, each bearing two protrud- houses and making of furniture.
ing pollen sacs. Seed cones terminal on short shoots
with spreading, 3–5 mm long, curved acicular leaves
nearly enclosing the receptacle only. Ripe receptacle Dacrycarpus vieillardii (Parl.) de Laub., J. Arnold
3–4 mm long, warty, reddish. Ripe seeds 1(–2) on a Arbor. 50: 326. 1969. Podocarpus vieillardii Parl.,
receptacle, subglobose, 4–6 mm long, including the in Candolle, Prodr. 16 (2): 521. 1868. Type: New
smooth, light yellowish brown to blackish brown, Caledonia: Grande Terre, Province Sud, near Poila,
sometimes glaucous epimatium, with a longitudinal E. Vieillard 1262 (holotype P).
grooved crest terminating in a curved protruding
apex.
Etymology
The species epithet commemorates the French bota-
Distribution
nist Eugène Vieillard (1819–1896).
Malesia: Borneo [Kalimantan, Balikpapan (ex-
tinct?)] Sulawesi (Latimodjong Mts.), Philippines
Vernacular names
(Luzon); Papuasia: New Guinea.
TDWG codes: 42 BOR-KA PHI SUL 43 NWG-IJ No vernacular names are recorded for this species.
NWG-PN
Description
Ecology
Shrubby to large trees to 25 m tall (usually much
Dacrycarpus steupii is common in mossy forest smaller); trunk up to 50 cm d.b.h.; crown open,
and subalpine shrubberies from montane to alpine often candelabra-shaped. Bark exfoliating in short
zones. Depending on the vegetation type it is a shrub strips, dark brown weathering greyish white. Foliage
or a tall emergent tree; its best growth is in protected branches with leaves of two kinds, acicular and long-
gullies in the upper montane forest where trees can er flattened. Leaves on primary or secondary shoots
grow taller. In peaty, wet tussock grasslands at high and fertile shoots spirally arranged, ­decurrent,
 subulate-acicular, curved inward at tip, appressed
Ecology
to imbricate or slightly spreading on lateral foliage
branches, keeled abaxially, 2–7 × 0.6–1 mm, api- Dacrycarpus vieillardii is common on and probably
culate. Leaves on many 1–5 cm long terminal and restricted to the serpentine ultramafic rocks and la-
deciduous branchlets of younger trees bilaterally teritic soils derived from these. In moist draws and
flattened, distichous, decurrent, slightly S-curved, along river banks it grows to a small tree up to 10
with parallel smooth margins, (3–)7–10(–12) mm m, only in forest can it attain taller stature. It is tol-
long (shortest at base and apex of branchlet), 1–1.2 erant of flooding and thereby avoids competition
mm wide; apex curved forward, apiculate; this type from most other trees. It has been found from near
of branchlets is replaced on larger trees by those sea level to about 900 m altitude. In so-called tall
328 with shorter, less distichous, bilaterally flattened ‘maquis minier’ it is often accompanied by Casuari-
leaves intermediate in size and shape. Stomata on naceae, Myrtaceae (e.g. Melaleuca) and conifers like
all sides of leaves (leaves amphistomatic) in a few Dacrydium araucarioides; in taller rain forest it can
intermittent rows on each face from base to apex; grow with Agathis lanceolata and other Myrtaceae
leaves often glaucous. Pollen cones lateral on foli- (Metrosideros, Weinmannia).
age branchlets on very short shoots, subtended by
a few small leaves, short when immature, at matu-
Conservation
rity elongating to 7–10 mm long and 1 mm wide.
Microsporophylls with triangular, apiculate apex, IUCN: LC
ca. 0.6 × 0.4 mm, each bearing two protruding pol-
len sacs. Seed cones terminal on short shoots with
Uses
spreading, 1–2 mm long leaves up to the receptacle.
Ripe receptacle 2–3 mm long, warty, green turning There are few large trees of this species and most
purplish, with one protruding green bract leaf. Ripe individuals occur along streams in ultramafic soil,
seeds 1(–2) on a receptacle, subglobose to oval, 5–6 × which dwarves them. Consequently, there is no
4 mm, including the smooth, grey-white or glaucous commercial exploitation of the timber of this species
epimatium, with an obtuse double apex from one of in New Caledonia.
which descends a small ridge on either side.

Distribution
New Caledonia: Grande Terre.
TDWG codes: 60 NWC
 figure 62. Callitris columellaris in Kings Canyon N.P., Australia

figure 60. Austrotaxus

spicata tree in New Caledonia
(photo M. Gardner)
figure 61. Callitris
­canescens in Western Australia

figure 64. Callitris

­macleayana seed cones

figure 65. Callitris muelleri in New South Wales, Australia

figure 63. Callitris

­macleayana trees in the
­Herberton Range, Queensland

figure 66. Callitris

­muelleri seed cones
f i gu re 67. Callitris preissii at Woodman Point, Western Australia
f i g u re 6 8 . Callitris rhomboidea seed cones,
Grampian Mts., Victoria, Australia
f i g ure 6 9. Callitris roei in Fitzgerald
River N.P., Western Australia

f i g u re 7 0. Callitris roei seed cones and foliage

fig ure 7 3. Calocedrus formosana

­foliage and seed cones
f i gu re 7 1 . Calocedrus decurrens tree in
figure 74. Calocedrus
the Sierra Nevada, California, USA
­macrolepis foliage and pollen cones
→

figure 72. Calocedrus

­decurrens trunk in the
Sierra Nevada

figure 7 5. Calocedrus rupestris foliage

and pollen cones (photo L. Aveyanov)
figure 7 6. ­Cathaya
­argyrophylla tree
in ­Sichuan (photo
H. ­Nimsch)

figure 7 7. Cathaya
­argyrophylla seed cones
(photo S. X. Yu)
figure 7 8. Cedrus
atlantica pollen cones

figure 79. Cedrus

deodara seed cones

figure 80. Cedrus libani var. libani in the Taurus Mts., Turkey

figure 81. Cephalotaxus

­fortunei var. fortunei
pollen cones and ­foliage

figure 83. Cephalotaxus

­harringtonii var. harring­
tonii pollen cones

figure 82. ­Cephalotaxus

­fortunei var. fortunei
ripe seeds
 fig ure 84. Cephalotaxus
harringtonii var. harringtonii
ripe seeds
figure 87. Chamaecyparis
formosensis foliage
and seed cones

f i g u re 85. Cephalotaxus mannii

leaves and seeds (photo L. Averyanov)

figure 86. Chamaecyparis formosensis and C. obtusa in Chilan Shan, Taiwan

f i g u re 88. Chamaecyparis lawsoniana

pollen cones

f i g u re 8 9. Chamaecyparis
lawsoniana seed cones

figure 91. Cryptomeria japonica seed cones

fi g u re 9 0. Chamaecyparis
thyoides var. thyoides trunk in
North Carolina, USA
fi g u re 92 . Cunninghamia konishii ­foliage

figure 95. Cupressus

­dupreziana tree in the Sahara
(Tassili n’Ajjer, Algeria)

figure 93. Cunninghamia

­lanceolata seed cones
figure 96. Cupressus
­guadalupensis var. forbesii
in California, USA

figure 94. Cupressus ari-

zonica var. arizonica seed cones

figure 98. Cupressus macrocarpa near Monterey, California, USA

figure 99. Cupressus ­torulosa
var. torulosa
seed cones

fig ure 97. Cupressus

­lusitanica var. benthamii
seed cones
 figure 100. Dacry-
carpus cinctus foliage and
cones (photo T. Utteridge)

figure 101. Dacry­

carpus dacrydioides tree in
North Island, New Zealand

figure 103. Dacrycarpus dacrydioides

seed cones
figure 104. Dacrycarpus imbricatus var.
imbricatus flushing foliage

figure 102. Dacrycarpus

dacrydioides trunk

figure 10 6. Dacrycarpus kinabaluensis

seed cones
fi g u re 1 05 . Dacrycarpus kinabaluensis tree on Mt Kinabalu, Borneo
 figure 108. Dacrydium araucarioides
foliage and pollen cones
figure 107. Dacrydium araucarioides
in New Caledonia

f i g u re 1 0 9 . Dacrydium beccarii
foliage with pollen cones

fi gu re 1 1 1 . Dacrydium comosum
at Gunung Ulu Kali, Malaysia
figure 110. Dacrydium comosum
canopy at Gunung Ulu Kali

fig ure 112. Dacrydium

­comosum foliage

figure 113. Dacrydium

­cupressinum in North Island,
New Zealand
 figure 116. Dacrydium gracile
at Bukit Tupai, Mt Kinabalu

figure 117. Dacrydium guillauminii

fi g u re 1 1 4 . Dacrydium elatum at Gunung Ledang, Malaysia in New Caledonia (photo A. Schmidt)

fi g u re 1 15 . Dacrydium gibbsiae small tree on Mt Kinabalu, Borneo

fig ure 118. Dacrydium xanthandrum tree in the

Crocker Range, Borneo

figure 119. Dacrydium xanthandrum

trunk in the Crocker Range

figure 12 0. Diselma archeri in Cradle

Mountain N.P., Tasmania, Australia
Dacrydium Sol. ex G. Forst., Pl. Escul. Ins. Ocean. Austr. Comm. Bot.: 80. 1786.
[Fl. Ins. Austr. Prod.: 92. 1786] Type: Dacrydium cupressinum Sol. ex G. Forst.
(Podocarpaceae).

Metadacrydium Baum.-Bod., Syst. Fl. Neu-Caledo-

Distribution
nien 5: 76. 1989. Type: Metadacrydium araucarioides
(Brongn. & Gris) Baum.-Bod. [Dacrydium araucari- Continental SE Asia: S China (Hainan); Indochina
oides Brongn. & Gris] (excluding Myanmar). Malesia: from Peninsular
Malaysia to the D’Entrecasteaux Islands (PNG), but
Greek: dakryon = a tear; referring to drops of resin excluding Jawa & Lesser Sunda Islands. SW Pacific: 337
exuded by the tree. Solomon Islands; New Caledonia; Fiji; New Zea-
land.
Description
Taxonomic notes
Dioecious or rarely monoecious evergreen shrubs
or trees. Resin in bark and (1 canal) in leaves. Bark The genus Dacrydium is one of the two earliest de-
hard, with numerous small lenticels, becoming scribed genera in the family Podocarpaceae (the oth-
fissured and scaly, exfoliating in plates. Primary er is Podocarpus) and it has in the past accomodated
branches in pseudo-whorls, numerous or sparse, numerous species, which are at present classified
spreading and assurgent, repeating the same pattern in other, in part segregate, genera, leaving a genus
(Rauh’s model), or with spreading and/or pendulous Dacrydium with a narrower, more precise circum-
foliage branches; latter foliage branches not termi- scription and at present containing 22 species. The
nating in distinct buds. Leaves spirally arranged, di- segregate genera are, in order of publication date,
morphic, scale-like and subulate or acicular, usually Lepidothamnus (1860), Acmopyle (1903), Falcati­
curved inwards, gradually transitional from juvenile folium (1969), Parasitaxus (1972), Halocarpus (1982),
to adult forms, keeled, triangular or quadrangular Lagarostrobos (1982) and, segregated again from
in cross-section, sometimes dorsiventrally flattened Lagarostrobos, Manoao (1995). The circumscription
(i.e. wider than thick). Stomata on all sides (leaves of Dacrydium has remained open to various inter-
amphistomatic). Pollen cones solitary or in groups, pretations until quite recently (Quinn, 1982), and as
terminal or lateral on foliage shoots, axillary on a result conifer species that do not properly belong
short scaly peduncles or nearly sessile, becoming in the genus (even if including its segregates) or not
short cylindrical; microsporophylls peltate, triangu- even in Podocarpaceae, have been ascribed to it. An
lar to apiculate, with two basal pollen sacs contain- informal classification into sections A, B and C by
ing bisaccate pollen. Seed cones terminal on (short) Florin (1931) divided the genus in its wider sense and
foliage branchlets, small, consisting of several spiral- only the species of his section B are still included in
ly arranged, scale-like or leaf-like (acicular) bracts Dacrydium. In subsequent accounts, when the genus
following smaller scales; fertile bracts with a single was circumscribed in a narrower sense, no attempts
more or less inverted ovule on the adaxial side; the were made to subdivide it (e.g. De Laubenfels, 1969,
whole enlarged, swollen and red when ripe forming 1988; Gaussen, 1974; Quinn, 1982). No phylogenetic
a small receptacle with protruding bract tips in most analysis of the genus has been published; at best a
species. Seeds 1–2(–3) per cone, becoming more or few species were included in investigations into the
less erect and usually standing obliquely from axis of phylogeny of the family Podocarpaceae at the genus
subtending receptacle, protruding from a cup-like level. There is therefore no sound basis upon which
epimatium covering its basal part only and ripening to base a classification of Dacrydium. Divisions of
to dark lustrous brown or nearly black. the genus based on a pre-selected few morphologi-
cal characters would, as is explained under Podocar-
22 species. pus, most likely be artificial. It is for this reason that
this relatively large (in conifer terms) genus must be
treated here in geographical groupings for the sake of
constructing identification keys that are ­manageable
 and may work in the field. Few species are in culti-
Key to the species of Dacrydium in Borneo
vation outside their native areas, so this should not
create undue problems. Two islands stand out as Juvenile leaf forms gradually merge into adult leaf
logical entities for this purpose: New Caledonia and forms and the ‘extremes’ of both types should there-
Borneo. The five species in New Caledonia are, as fore be taken for comparison with the character
the other conifers there, endemic. Most of the seven states used in this key. Adult leaf forms are in most
species of Borneo are also restricted to the island, species shorter than juvenile leaf forms.
but three species are also found elsewhere. These,
and the remainder (11 species) are dealt with in the 1a. Adult leaves scale-like, appressed, 1–1.5 mm
third key. long D. elatum
338 1b. Adult leaves acicular, linear or linear-lanceo-
late, spreading, (1.5–)3–10(–13) mm long 2
Key to the species of Dacrydium in New
2a. Adult leaves and juvenile leaves similar, ac-
Caledonia
icular, spreading wide to nearly 90°, nearly
Juvenile leaf forms gradually merge into adult leaf straight, 5–10 mm long, 0.7–1 mm wide, flat-
forms and the ‘extremes’ of both types should there- tened D. ericoides
fore be taken for comparison with the character 2b. Adult leaves and juvenile leaves of different
states used in this key. Adult leaf forms are in most length (adult leaves much shorter); adult leaves
species shorter than juvenile leaf forms. spreading forward or in one species wide to
90° 3
1a. Shrubs or dwarfed trees up to 2.5 m (rarely 3a. Adult leaves spreading wide to nearly 90°,
to 4 m) tall. Juvenile and adult leaves similar curved forward or nearly straight. Microsporo-
in size and shape or adult leaves only slightly phylls of pollen cones with a rostrate apex
shorter, the shortest at least 5 × longer than D. xanthandrum
wide 2 3b. Adult leaves spreading forward more or less
1b. Small or large trees. Juvenile and adult leaves close to the shoot, usually curved. Microsporo-
very different in size and shape, the shortest less phylls of pollen cones triangular or in one spe-
than 4 × longer than wide 3 cies lanceolate 4
2a. Adult leaves slightly shorter than juvenile 4a. Adult leaves very slender and soft, 0.3–0.4 mm
leaves, 5–10 mm long (juvenile leaves 10–15 mm wide, slightly curved or nearly straight
long), imbricate but with a free apex, narrowly D. beccarii
lanceolate. Seeds 3–4 × 2 mm D. ×suprinii 4b. Adult leaves more or less rigid, (0.4–)0.5–1.2
2b. Adult leaves equally long as juvenile leaves, mm wide, curved at least towards the apex 5
(10–)13–15(–17) mm long, spreading but curved 5a. Adult leaves robust, 0.9–1.2 mm wide, linear or
forward, acicular. Seeds 4–5 × 2.5 mm linear-lanceolate. Pollen cones 20–25 × 4.5–7
D. guillauminii mm; microsporophylls lanceolate, 4–5 mm
3a. Adult leaves linear-lanceolate, 0.6–0.8 mm long D. gibbsiae
wide; apex pungent. Pollen cones 4–7 mm long, 5b. Adult leaves slender, 0.4–0.8 mm wide, acicular
1.2 mm wide D. lycopodioides or linear. Pollen cones 6–12 × 2 mm; microspo-
3b. Adult leaves oblong, incurved apically, 1–1.8 rophylls triangular-apiculate, 1–1.5 mm long 6
mm wide; apex obtuse. Pollen cones 8–18 mm 6a. Adult leaves linear, nearly as thick as wide,
long, 1.5–4 mm wide 4 0.4–0.8 mm wide, keeled sharply on the abaxial
4a. Adult leaves imbricate, strongly incurved api- side; apex obtuse or short apiculate. Seed cones
cally hiding the apex from view. Pollen cones terminal on foliage shoots; epimatium covering
3–4 mm wide. Seed cones 8–10 mm long, with the basal third of the seed D. pectinatum
1–3 seeds D. araucarioides 6b. Adult leaves acicular, wider than thick (trans-
4b. Adult leaves spreading 30–45° from shoot; apex verse-triangular in cross-section), 0.4–0.5
incurved but free and visible. Pollen cones 1.5–2 (–0.7) mm wide, keeled weakly on the abaxial
mm wide. Seed cones 2–3 mm long, with a sin- side; apex apiculate. Seed cones lateral on foli-
gle, rarely 2 seeds D. balansae age shoots; epimatium covering the basal half
of the seed D. gracile
 7b. Juvenile leaves 5–20(–25) mm long, 0.2–0.6
Key to the species of Dacrydium in regions
mm wide, not flattened; adult leaves not flat-
outside Borneo and New Caledonia
tened 8
Juvenile leaf forms gradually merge into adult leaf 8a. Bracts subtending seed cones longer than the
forms and the ‘extremes’ of both types should there- leaves immediately below these, 3–8 mm long,
fore be taken for comparison with the character completely hiding the cones 9
states used in this key. Adult leaf forms are in most 8b. Bracts subtending seed cones shorter or equally
species shorter than juvenile leaf forms. long as the leaves immediately below these, 1–3
mm long, exposing the cones and/or the epi-
1a. Adult leaves 1–2(–3) mm long, scale-like or ac- matium 12
icular, appressed or spreading forward and in- 9a. Juvenile leaves 8–14 mm long, strongly curved; 339
curved, 0.2–0.5(–0.7) mm wide 2 apex apiculate. Pollen cones 8–12 × 1.8–2 mm;
1b. Adult leaves (1.5–)3–10(–30) mm long, acicu- microsporophylls triangular
lar or linear-lanceolate, spreading forward or D. cornwallianum
outward, if incurved mostly longer than 3 mm, 9b. Juvenile leaves 10–20 mm long, straight or
0.3–1.2 mm wide 5 slightly curved; apex pungent. Pollen cones (7–)
2a. Adult leaves scale-like, appressed, on slender, 10–18 × 2–2.5 mm; microsporophylls apiculate
cord-like shoots 3 or elongated 10
2b. Adult leaves acicular, spreading forward and 10a. Juvenile leaves very slender, 0.2–0.3 mm wide,
incurved, on slender, brush-like shoots 4 not keeled abaxially. Microsporophylls of pol-
3a. Juvenile leaves 5–12 mm long, 0.4–0.6 mm len cones short, apiculate; seeds 3.5–4 mm long,
wide, keeled abaxially; bracts at base of seed lustrous brown D. nidulum
cones 3 mm long D. novoguineense 10b. Juvenile leaves 0.4–0.6 mm wide, keeled abaxi-
3b. Juvenile leaves 10–15 mm long, 0.3 mm wide, ally. Microsporophylls of pollen cones elon-
keeled on 4 sides; bracts at base of seed cones gated, with incurved apex; seeds 4–5 mm long,
1 × 1 mm, triangular D. elatum brown or black 11
4a. Adult leaves sharply keeled abaxially; apex ob- 11a. Adult leaves partly imbricate, straight with in-
tuse or sometimes acute D. nausoriense curved apex, 0.5–0.6 mm wide. Pollen cones
4b. Adult leaves not keeled but transverse-triangu- 7–9 × 2.5 mm. Bract scales subtending seed
lar in cross-section; apex acute-pungent to acu- cones 3–5 mm long D. medium
minate D. leptophyllum 11b. Adult leaves free, spreading forward, incurved,
5a. Juvenile leaves 15–33 mm long, 0.8–1 mm wide; 0.3–0.4 mm wide. Pollen cones 10–16 × 2–2.5
adult leaves (5–)7–25(–30) mm long, 0.5–1.2 mm. Bract scales subtending seed cones 6–8
mm wide 6 mm long D. magnum
5b. Juvenile leaves 5–20 mm long, 0.2–0.6(–1) mm 12a. Juvenile leaves 5–8 mm long, not much longer
wide; adult leaves (1.5–)2–10 mm long, 0.3–1 than adult leaves, keeled on 4 sides; adult leaves
mm wide 7 0.8–1 mm wide. Seed cones when fully devel-
6a. Shrubs or dwarf trees to 6 m tall; adult leaves oped 6–8(–10) mm long, swollen succulent and
crowded, spreading and curved forward, 12– orange to red; seeds lustrous black
25(–30) mm long, 0.6–1.2 mm wide D. cupressinum
D. comosum 12b. Juvenile leaves 10–20(–25) mm long, much
6b. Trees to 35 m tall, sometimes shrubs at high- longer than adult leaves, keeled abaxially; adult
est altitudes; adult leaves spreading at ca. 90°, leaves 0.3–0.8 mm wide. Seed cones minute, ca.
straight or curved, (5–)7–10(–13) mm long, 3 mm long; seeds lustrous brown 13
0.5–0.8 mm wide D. xanthandrum 13a. Adult leaves (4–)6–10 mm long, acicular, soft,
7a. Juvenile leaves 5–7 mm long, 1 mm wide, flat- 0.3–0.4 mm wide. Seeds 1–2(–3) per cone
tened, 0.2–0.3 mm thick; adult leaves 2–4 mm D. beccarii
long, 0.8–0.9 mm wide, flattened, 0.2–0.3 mm 13b. Adult leaves (1.5–)3–5(–7) mm long, linear, rig-
thick, linear-lanceolate D. spathoides id, 0.4–0.8 mm wide; Only 1 seed per cone
D. pectinatum
 Dacrydium araucarioides Brongn. & Gris, Ann.
Distribution
Sci. Nat. Bot., sér. 5, 6: 244. 1866. Metadacrydium
araucarioides (Brongn. & Gris) Baum.-Bod., New Caledonia: Grande Terre, Province Sud.
Syst. Fl. Neu-Caledonien 5: 76. 1989. Type: New TDWG codes: 60 NWC
Caledonia: Grande Terre, Province Sud, Canala,
[“montagnes au-dessus de Canala”], E. Vieillard
Ecology
1277 (lectotype P). Fig. 107, 108
This species is restricted in its occurrence to the
­ultramafic substrates derived from peridotite or ser-
Etymology
pentine; it is particularly abundant on laterite soils
340 The species epithet means ‘similar but not equal to and ‘ironstone’ (‘cuirasse de fer’) in the Plaine des
Araucaria’ and refers to the foliage branches and Lacs and the surrounding hills. It occurs in low, open
their leaves. woodland on mountain ridges and in tall ‘maquis
minier’ on slopes and plains, from just above sea
level to ca. 1000 m altitude. It is accompanied by nu-
Vernacular names
merous angiosperms; one of the most characteristic
This species does not have a common name. small trees looking very similar from a distance is
Gymnostoma chamaecyparis (Casuarinaceae). As-
sociated conifers are e.g. Araucaria muelleri, Aga-
Description
this ovata, Retrophyllum minus, and Neocallitropsis
Small trees 2–7 m tall; trunk to 15 cm d.b.h., erect, pancheri. This type of vegetation is prone to fires and
monopodial. Bark smooth becoming rough, with regeneration of the conifers and other small trees is
horizontal ridges, breaking into small scales, brown generally slow due to the extreme poverty of nutri-
weathering grey; inner bark slightly fibrous, red- ents in the soil.
brown. Primary branches few, spreading and assur-
gent, with foliage branches towards the end, forming
Conservation
a candelabra crown but often irregular and open.
Juvenile leaves on seedlings and saplings, acicular, IUCN: LC
straight on seedlings to curved on saplings, 7–12 mm
long, 0.7–1 mm wide and thick, keeled on four sides;
Uses
apex acute to apiculate. The (pen)ultimate branch-
lets with adult leaves 4–8 mm thick; the leaves spi- No uses are recorded of this species.
rally arranged, imbricate, spreading from base at
<30° and strongly incurved towards apex, oblong,
2–4(–5) × 1–1.5 mm, keeled from base abaxially, be- Dacrydium balansae Brongn. & Gris, Bull. Soc.
coming convex towards the obtuse (invisible) apex, Bot. France 16: 328. 1869. Metadacrydium balansae
lustrous green. Stomata on juvenile leaves on all (Brongn. & Gris) Baum.-Bod., Syst. Fl. Neu-Cale-
sides in lines between ribs (leaves amphistomatic), donien 5: 76. 1989. Type: New Caledonia: Grande
on adult leaves largely confined to the adaxial side Terre, Province Sud, Bourail, [“au-dessous de Téné
but a few near base abaxially. Pollen cones terminal près de Bourail”], B. Balansa 1380 (holotype P).
or lateral on very short branchlets, cylindrical, 9–18
× 3–4 mm; microsporophylls similar to adult leaves
Etymology
but smaller and shorter, nearly triangular with in-
curved apex; pollen sacs not visible due to imbricate The species epithet commemorates the plant collec-
microsporophylls. Seed cones terminal, subtended tor Benjamin Balansa (1825–1892), who collected in
by 3 mm long leaves; bracts elongated to ca. 5 mm, New Caledonia in the years 1868–1872.
less curved than the leaves, covering the 8–10 mm
long cone up to the seed, turning purplish red at ma-
Vernacular names
turity; epimatium at base of seed. Seeds single or 2–3
per cone, partly covered by the bracts, ovoid with a No vernacular name is known for this species.
slightly constricted apex, 4–4.5 mm long, brown.
 but identity of such trees observed lacks evidence
Description
from herbarium specimens. It is not uncommon in
Trees 3–12(–20?) m tall; trunk to 30 cm d.b.h., erect, some localities, but never forms large populations.
monopodial. Bark becoming rough, with horizon- It is associated with several other conifers (Agathis
tal ridges, breaking into irregular scales, brown ovata, Araucaria spp., Dacrydium araucarioides)
weathering grey; inner bark fibrous, red-brown. and numerous angiosperms.
Primary branches spreading and assurgent, with
numerous foliage branches towards end, forming a
Conservation
domed crown but often irregular and open. Juvenile
leaves on seedlings and saplings, acicular, straight IUCN: LC
on seedlings to curved on saplings, 9–13 mm long, 341
0.7–1 mm wide and thick, keeled on four sides; apex
Uses
acute to apiculate. Adult leaves spreading 30–45°
from shoot and free, length variable on branchlets The commonly small stature of this species makes
with shorter leaves often at beginning and end of it an unlikely source of timber despite good wood
branchlet, 2–5(–7) × 1–1.8 mm, oblong but widest properties; an occasional larger specimen may have
near base, curved forward, keeled abaxially, flat or been taken for this purpose. It is not known in cul-
concave adaxially; apex incurved, obtuse. Stomata tivation.
on juvenile leaves on all sides in several intermittent
lines (leaves amphistomatic), on adult leaves idem
but not extending to apex on the abaxial side. Pol- Dacrydium beccarii Parl., in Candolle, Prodr. 16
len cones terminal or lateral on very short branch- (2): 494. 1868. Type: Malaysia: Sarawak, Gunung
lets, cylindrical, becoming curved when expanded Pueh (Mt. Poe), O. Beccari 2385 (lectotype FI). Pl.
to 8–15 × 1.5–2 mm; microsporophylls similar to 13, Fig. 109
adult leaves but smaller and shorter, nearly triangu-
lar with incurved apex; pollen sacs not visible due
Etymology
to imbricate microsporophylls. Seed cones termi-
nal, 2–3 mm long, subtended by 2 mm long leaves; This species was named after the Italian botanist
bracts elongated to 3.5–4 mm, curved at apex, cover- Odoardo Beccari (1843–1920), who collected the
ing the cone up to the seed, turning purplish red at type specimen.
maturity; epimatium basal to seed. Seeds single or
rarely two per cone, obliquely oriented, often partly
Vernacular names
covered by bracts, ovoid with a slightly constricted
apex, 4.5–5 × 3.5 mm, brown. Many vernacular names are in use for this species,
differing from island to island and even locally.
Some are: kayu embun (Borneo), ekor kuda (Ma-
Distribution
laya), netukuria (New Guinea).
New Caledonia: Grande Terre.
TDWG codes: 60 NWC
Description
Shrubs or trees of medium size, to 25 m tall; trunk to
Ecology
80 cm d.b.h., much branched eventually developing
Dacrydium balansae is a small tree occurring mostly a broadly spreading dense crown, sometimes more
on ultramafic soils derived from serpentine at ele- or less flat-topped. Bark scaly; outer bark brown;
vations from near sea level to 800–900 m. Its stat- inner bark finely fibrous, exuding red resin from
ure is partly dependent on soil and exposure, with slash. Foliage of two indistinct types, with juvenile
dwarfed trees predominantly on the most metallif- leaves and adult leaves mainly differing in length
erous rocks with very thin soil that support ‘maquis and width. Juvenile leaves spreading at 30–70° from
minier’ vegetation and taller trees occurring infre- shoot, acicular, very slender, 15–20(–25) × 0.3–0.6
quently in moist ravines that support taller forest. mm, usually curved forward, keeled abaxially, pun-
Here individuals rarely may attain hights near 20 m, gent. Adult leaves densely set, spreading at 30–45°
342

Pl ate 13. Dacrydium beccarii. 1. Habit of tree. 2. Branch with foliage. 3. Leaf. 4. Branchlet with juvenile
leaves. 5. Branchlet with leaves and seed cones. 6. Seed cone with seed.
forward, sometimes closer to shoot, acicular, very
Uses
slender and soft, slightly curved or nearly straight,
(4–)6–10 × 0.3–0.4 mm, triangular in cross-section Dacrydium beccarii is locally exploited for timber;
and slightly wider than thick, keeled abaxially, pun- the wood is used in building houses and for mak-
gent, dark green, flushing light green. Stomata on ing drums (New Guinea). The bark is used in some
all sides (leaves amphistomatic), in 1–2 inter­mittent villages in the highlands as an insulation material in
lines on each face to apex. Pollen cones lateral or the walls of round small houses or huts.
terminal on branchlets with adult leaves (sometimes
with juvenile leaves), 7–10 × 2.5–3 mm at maturity;
microsporophylls triangular, 1 mm wide at base, Dacrydium comosum Corner, Gard. Bull. Straits
with extended narrow apex 1 mm long and 0.3 mm Settlem. 10: 244, t. 10. 1939. Type: Malaysia: Malaya, 343
wide at base, turning brown; 2 basal reddish pol- Negeri Pahang, Pine Tree Hill, E. J. H. Corner SFN
len sacs. Seed cones terminal on leaved branchlets, 33222 (holotype SING). Fig. 110, 111, 112
subtended by yellowish involucral leaves, 3 mm
long, covered with triangular bract scales 1 × 2 mm,
Etymology
slightly swelling and turning red at maturity. Seeds
single or 2, rarely 3, obliquely positioned, 3.5–4 mm The species epithet means ‘bearing a tuft of leaves’
long, covered for the basal third by the epimatium (Latin comosus = ‘with much or long hair’).
and turning lustrous dark brown.
Vernacular names
Distribution
No common names have been recorded for this spe-
Malesia: Borneo, Peninsular Malaysia, Maluku cies.
[Moluccas], Philippines, Sulawesi, Sumatera; Papua-
sia: New Britain, New Guinea, Solomon Islands.
Description
TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
MLY-PM MOL PHI SUL SUM 43 BIS NWG-IJ NWG-PN Small trees to 15 m tall, with a single stem to 50 cm
SOL-SO d.b.h., on exposed ridges dwarfed and flat-topped.
Branches spreading and ascending in candelabra
fashion, making an umbrella-shaped crown with
Ecology
crowded, upturned foliage branches at ends of pri-
Dacrydium beccarii is most common in low canopy mary branches. Juvenile leaves and adult leaves very
(to 20 m) mossy forest on leached, podzolic sandy similar. Juvenile leaves spreading more perpen-
soils (kerangas), where it can dominate the canopy dicular to branch, acicular, curved forward, very
especially on ridges of mountains at altitudes from narrowly tapering, to 33 mm long, ca. 1 mm wide
500 m to 2500 m a.s.l. It has also been found on karst at base, keeled abaxially, ending in a pungent apex.
limestone, granite, and andesite, as well as in wet Adult leaves densely crowded and covering the shoot
peaty soil. In exposed locations on mountain sum- completely, spreading forward, acicular, straight
mits it becomes an emergent low tree above dense or slightly curving outward, 12–25(–30) mm long,
scrub. It is associated with other conifers, such as 0.6–1.2 mm wide at or just above a slightly widened
Agathis spp., Dacrycarpus spp., Falcatifolium ­gruezoi base, flattened with a sharp abaxial keel, gradually
(e.g. on Mt. Halcon in the Philippines), Nageia wal- tapering to a pungent apex. Stomata on both sides
lichiana, and sometimes Sundacarpus amarus. On (leaves amphistomatic) in a few intermittent lines
mountain ridges it can better compete with the nu- nearly to apex. Pollen cones mostly lateral on foliage
merous angiosperms in these forests, where these branches, subtended by a few short leaves ca. 5 mm
angiosperms remain smaller trees and shrubs. long, 8–10 × 3 mm; microsporophylls terminating
in a narrow extension, 1.5–2 mm long and 0.5 mm
wide at base, with two basal pollen sacs. Seed cones
Conservation
few, terminal on short foliage branchlets, ca. 3 mm
IUCN: LC long, nearly hidden by leaves, with several lanceo-
late, coloured bracts and 1(–2) exposed, ovoid, light
 brown seeds 4–5 mm long covered at base by the
Vernacular names
­epimatium.
No vernacular names are known for this species.
Distribution
Description
Peninsular Malaysia (Genting Highlands, Gunung
Hulu Kali, Negeri Pahang). Trees to 30 m, monopodial, erect; trunk d.b.h. to
TDWG codes: 42 MLY-PM 50 cm; crown elongated, dense, often with fastig-
iate branching. Bark thin and hard, scaly and shed-
ding small flakes, brown, weathering grey; inner
Ecology
344 bark slightly fibrous and pinkish. Foliage in much
Dacrydium comosum occurs on exposed mountain branched tufts, spreading or assurgent in mature
ridges as a local dominant in stunted mossy forest trees. Leaves on seedlings and juvenile trees acicular,
on rocky acidic soil or shallow peat; in one area at 8–14 mm long, 0.4–0.5 mm wide and 0.2–0.3 mm
altitudes between 1170 and 1440 m a.s.l. (the alti- thick, spreading and strongly curved forward or
tudinal range is incompletely known). Occasional incurved, keeled abaxially, sharply apiculate. Adult
individuals may grow in forest below the ridge and leaves shorter and mostly uniform on branchlets,
attain tree size to reach the canopy. spirally arranged and decurrent at base, (2–)3–5(–8)
mm long, 0.6–0.8 mm wide and 0.3–0.4 mm thick,
keeled sharply on the abaxial side, spreading at 30°
Conservation
but strongly curved forward and with an incurved,
Dacrydium comosum is known from two separate apiculate apex. Intermediate forms of leaves occur
areas and in total from less than five localities within on younger trees. Stomata on all sides (leaves am-
these areas. Corner (op. cit.) mentioned four trees phistomatic), but extending to apex only on the
from the type locality, but the numbers at other lo- adaxial side where they are separated by a midvein.
calities are not known. There is a risk of fire threat- Pollen cones terminal or sometimes lateral, 8–12
ening these small populations coming from burning ×1.8–2 mm; microsporophylls triangular, 1–1.2 × 0.7
and clearing forest at lower elevations. Seed produc- mm, incurved, faintly keeled abaxially, obtuse, with
tion seems to be low and seedlings are sparse. two basal pollen sacs. Seed cones terminal, ca. 3 mm
IUCN: EN (B1a,b+B2a,b) long, subtended by short (1.5–2 mm) leaves followed
by longer (3–5 mm) cone bracts surrounding the
epimatium, becoming swollen and red at base. Seed
Uses
single, epimatium covering only the basal third or
No uses have been recorded of this species. less of the 4–5 mm long, lustrous brown seed.

Taxonomic notes
Dacrydium cornwallianum de Laub., Fl. Malesiana,
ser. 1, 10 (3): 366. 1988, [“cornwalliana”]. Type: Indo- This taxon was elevated to a species by De Lauben-
nesia: Papua, Danau Paniai [Wissel] Lakes, Arupa, fels in 1988 (op. cit.) from Dacrydium nidulum var.
C. Versteegh BW 3041 (holotype A, isotype K). araucarioides de Laub. (1969, op. cit.). It appears
to have sufficient distinct characters to merit this
Dacrydium nidulum de Laub. var. araucarioides de move, which is here accepted. The author based the
Laub., J. Arnold Arbor. 50: 293. 1969. Type: Indo- two taxa on different type specimens (duplicates) of
nesia: Papua, Danau Paniai [Wissel] Lakes, Arupa, the same collection taken from a single tree, lodged
C. Versteegh BW 3041 (holotype L). in different herbaria; he also described D. corn­
wallianum explicitly as a new species, yet cited the
earlier published variety as a synonym. The binomial
Etymology
D. araucarioides could not be used as it had already
Although no explanation was given, the species epi- been applied to a species from New Caledonia.
thet means ‘of Cornwall’.
 rough and scaly, peeling in elongated, thick flakes,
Distribution
light brown weathering grey. Young trees pyrami-
New Guinea. dal, crown of mature trees becoming domed, with
TDWG codes: 43 NWG-IJ NWG-PN spreading to ascending first order branches. Foliage
branches drooping to pendulous, slender. Juvenile
and adult leaves similar in shape but adult leaves
Ecology
shorter, especially on fertile branches. Juvenile leaves
This species is a dominant in mixed forest, or forms on seedlings to saplings and young trees, spreading,
nearly pure stands in swamp forest, and perhaps straight, soft and pliable, 5–8 mm long, 0.3–0.5 mm
also in mossy forest, between 1450 and 2300 m al- wide in seedlings and saplings, becoming stiffer and
titude. A photograph in Flora Malesiana 1, 10: 366 thicker on larger young trees, keeled on four sides, 345
(1988) shows a dense stand of this tree in the ecotone pungent, green but often turning bronze in winter.
between montane rainforest and swamp “on black Adult leaves 2–3(–4) mm long, 0.8–1 mm wide, de-
peat”. current, widest just above base where they spread
at 45–60°, straight or slightly curved, triangular in
cross-section, acute to acuminate. Leaves amphi­
Conservation
stomatic, but with most stomata on adaxial side in
Although apparently rare and only known from a many lines to apex, on abaxial side a few lines near
handful of collections and localities, this species oc- base (on adult leaves nearly absent on that side).
curs in habitat that is far removed from deforesta- Pollen cones terminal, ovate-oblong, 7–10 mm long,
tion or other land uses detrimental to the natural 4 mm wide; microsporophylls with elongated apex,
vegetation. It may be more common than the few 2–2.5 mm long, green, each with two yellow pollen
collections known seem to suggest. This species sacs at base. Seed cones terminal on short, curved
does not appear to occur within protected areas. branchlets or lateral, 6–8(–10) mm long; bracts
IUCN: LC short, scarcely protruding when the swollen, or-
ange to red cone is mature. Seeds solitary, obliquely
placed, the basal ⅓ covered by a green, later bright
Uses
red epimatium, seed ovoid, slightly flattened, 4–5
The timber of this species may be locally used for mm long, ripening to lustrous black.
construction of traditional houses in the villages of
the highland areas in western New Guinea.
Distribution
New Zealand, distributed widely on all main islands.
Dacrydium cupressinum Sol. ex G. Forst., Pl. Escul. TDWG codes: 51 NZN NZS
Ins. Ocean. Austr. Comm. Bot.: 80. 1786. Type not
designated (New Zealand, prob. coll. D. Solander
Ecology
on Cook’s first voyage). Fig. 113
Dacrydium cupressinum is a dominant or codomi-
nant emergent in mixed conifer (podocarp) forest
Etymology
or conifer-angiosperm forest at low to middle eleva-
The species epithet denotes a similarity with cy- tions up to 700 m a.s.l. These forests are (warm) tem-
presses (Cupressus). perate evergreen rain forests with year-round high
precipitation and are multi-layered with emergent
conifers, a canopy of conifers and/or angiosperms,
Vernacular names
an understorey of shrubs, tree ferns and palms, and
Red pine; rimu (Maori) a ground cover of ferns and mosses. In the kauri for-
est of Northland D. cupressinum is a minor compo-
nent, but in other forest types it is often the most
Description
common species, sometimes occurring in nearly
Large trees to 50(–60) m tall; trunk d.b.h. to 1.5–2 m, pure stands. Very large specimens may be around
erect, usually monopodial, sometimes forked. Bark 1000 years old, but few of these survive today in
 only remnants of a once extensive native lowland The most decoratively figured pieces were reserved
forest cover. Common associated conifer species for cabinet making. Due to severe depletion of this
are Agathis australis (restricted to the northern part natural resource, exploitation has been halted and
of the range of D. cupressinum), Podocarpus totara, this species is now protected from logging by law.
Prumnopitys ferruginea, P. taxifolia, Dacrycarpus As a slow growing tree it is not considered suitable
dacrydioides, Manoao colensoi, Phyllocladus tri- for plantation forestry and it finds only limited use
chomanoides and, less frequently, Halocarpus kirkii. as an amenity tree in gardens and parks, mainly in
Some common angiosperm trees are Beilschmiedia New Zealand.
tarairi (Lauraceae), Metrosideros spp. (Myrtaceae),
Weinmannia racemosa (Cunoniaceae) and Quin-
346 tinia acutifolia (Escalloniaceae), and there are many Dacrydium elatum (Roxb.) Wall. ex Hook., London
other genera and species. Dacrydium cupressinum is J. Bot. 2: 144, t. 2. 1843. Juniperus elata Roxb.,
a typical example of a long-lived ‘catastrophic regen- Fl. Ind. 3: 838. 1832. Type: Malaysia: Peninsular
eration’ conifer (Ogden & Stewart in Enright & Hill, Malaysia, Penang Island, Penang Hill, [“native
eds. 1995). of the Island of Pulo Pinang”], “Dr. Cantor” s.n.
(holotype K). Fig. 114
Conservation
Dacrydium pierrei Hickel, Bull. Soc. Dendrol. France
Dacrydium cupressinum is a widespread and com- 76: 74. 1930. Type: Vietnam: South Vietnam, Quan
mon species in New Zealand not considered in dan- Phu Quoc, L. Pierre 1396 (lectotype K, designated
ger of extinction at present. However, since nearly all here).
of the primary lowland forests in which it was abun-
dant have been cleared for agriculture, plantation
Etymology
forestry, and urban development within a period of
150 years ending around 1970, this (and other) tree The species epithet elatum means ‘tall’ and refers to
species has undergone a very severe reduction in its the large size some trees can attain.
area of occupancy (AOO) of at least 70% over “three
generations” (even though that reduction has now
Vernacular names
ceased) and would therefore qualify as Endangered
(EN) under IUCN criterion A1. The application of kayuru bukit, ru bukit (Malaya); ouk (Borneo);
this criterion to New Zealand trees remains con- sambinur (?)
troversial, because conservationists in that country
take the view that assessments of threat to species
Description
should only consider the present and future, not the
past. Trees of medium to large size, to 40 m tall; trunk
IUCN: LC to 1 m d.b.h., much branched eventually developing
a broadly spreading dense crown, sometimes more
or less flat-topped. Bark scaly; outer bark brown to
Uses
blackish; inner bark pink to red, exuding red resin
Rimu or Red pine was, when exploited, the major from slash. Foliage of two types, with juvenile leaves
timber tree in New Zealand as it was ubiquitous and with adult leaves. Juvenile leaves acicular, on
throughout the lowlands of the major islands and seedlings and saplings very slender, 10–15 × 0.3 mm,
attained large sizes. The following remark “… wood shorter and slightly thicker on young trees (especial-
close grained rather brittle and very durable… next ly at base of branchlets), straight or slightly curved
in estimation to kauri for house building.” [Mr forward, spreading at less than 45° from the shoot,
Clarke in Alan Cunningham’s New Zealand Her- keeled on four sides, pungent. Transitional leaves
barium, No. 332 (K)] indicates its importance. The sometimes present, but usually abrupt change to
wood is reddish brown in colour, often finely figured adult leaves. Adult leaves scale-like on slender, cord-
and very durable and strong. It was used in general like, much branching shoots up to 10 cm long and to
construction, for bridges, railway sleepers, fence 3 mm wide (ultimate branchlets usually 1 mm wide),
posts, indoor flooring and panelling, and furniture. appressed or rarely free at apex, rhombic to oblong-
rhombic, 1–1.5 × 0.3–0.5 mm on ultimate branchlets, forest and swamp or savannah D. elatum is usually
larger on leading shoots, keeled abaxially, obtuse growing along streams. It appears to be restricted to
or acute with incurved apex. Stomata on all sides acidic soils derived from granite or sandstone, often
(leaves amphistomatic), on juvenile leaves in narrow mixed with organic matter like peat.
grooves, on adult leaves in scattered lines, absent on
the distal part of the abaxial side, on the adaxial side
Conservation
to apex. Pollen cones terminal on branchlets with
adult leaves (sometimes with juvenile leaves), 4–8 IUCN: LC
× 1.2–1.5 mm at maturity, microsporophylls broad
triangular, 1 × 1 mm, acute-apiculate, with two ba-
Uses
sal reddish pollen sacs, turning brown. Seed cones 347
sometimes among juvenile foliage, but always ter- Dacrydium elatum is probably the main source of
minal on branchlets with adult leaves, 3–4 mm long, ‘sempilor’ wood in SE Asia, which includes spe-
covered with broadly triangular bract scales 1 × 1 cies of Dacrycarpus, Dacrydium, Falcatifolium and
mm, slightly swelling and turning red at maturity. Phyllo­cladus, between which the timber trade makes
Seeds single, obliquely positioned, 3.5–4.5 mm long, no distinction. It is a light and relatively hard wood
covered only at base by the epimatium and turning used in construction, window frames, doors, join-
lustrous black. ery, furniture, flooring, interior finishing, veneer,
and plywood as well as pulp for the paper industry.
A volatile oil resembling cedar oil can be distilled
Taxonomic notes
from the wood, which is done during the pulping
The morphological differences between Dacrydium process. This species is cultivated as an ornamental
pierrei and D. elatum as stated by Hickel (op. cit.) tree, mainly in countries within its natural range.
and Gaussen (1974) do not exist in the herbarium
specimens examined at K from ‘Indochina’. Among
these are 4 sheets of L. Pierre 1396 specifically men- Dacrydium ericoides de Laub., Fl. Malesiana,
tioned by Hickel for the alledged differences; one of ser. 1, 10 (3): 371. 1988. Type: Malaysia: Sarawak,
these sheets (Kew barcode K000288637) is here des- Merurong Plateau, Bukit Skelap, E. F. Brunig, SFN
ignated as the lectotype of D. pierrei Hickel. 8722 (holotype L).

Distribution Etymology
S China: Guangxi; Indochina: Kampuchea [Cam- The species epithet refers to a supposed similarity
bodia], Laos, Thailand, Vietnam; Malesia: Borneo with Ericaceae (Heather family) or the genus Erica.
(Sabah and Sarawak), Philippines (Sibuyan Is.),
Peninsular Malaysia (including Penang Island), Su-
Vernacular names
matera (W coast).
TDWG codes: 36 CHS-GX 41 CBD LAO THA VIE 42 sempilor (in the Bintulu dialect, given with the name
BOR-SB BOR-SR PHI MLY-PM SUM D. beccarii Parl. as the original identification of the
type collection)
Ecology
Description
Dacrydium elatum occurs in evergreen tropical rain-
forest from the lowlands up into the mountains at Trees 10–18 m tall; trunk to 30 cm d.b.h., erect and
altitudes between 100 m and 1350 m a.s.l. It is often single-stemmed, branching to form a spreading
found in wet places such as peat swamp forests in- crown. Bark scaly, brown; inner bark more or less
termixed with patches of kerangas forest. In the lat- fibrous, reddish. Foliage branches mostly droop-
ter type of vegetation, which occurs on nearly white ing or nearly pendulous, with more or less persist-
sandy soil, conifers often dominate in the forest, with ently juvenile type leaves becoming only slightly
Agathis, Dacrydium and Podocarpus usually present. shorter in the adult form on mature trees. Leaves
In more open situations in the border zone between linear, straight, spreading at almost right angles to
 shoot except on new growth, 5–10 mm long, 0.7–1 for logging, as the forest quality from that point of
mm wide, 0.2–0.3 mm thick, flattened but becoming view is poor on exposed mountain ridges. It is listed
slightly concave towards apex, sharply keeled on the as Vulnerable purely on the basis of its limited ex-
abaxial side, with a faint midrib adaxially, narrow- tend of occurrence (EOO); it is however possible
ing abrubtly at the distal end to an apiculate apex. that other localities will be found in future.
Stomata in two bands of 6–8 lines on adaxial side IUCN: VU (D2)
from base to apex, only a few near base abaxially
(leaves weakly amphistomatic). Pollen cones mostly
Uses
lateral on foliage branches, subtended by reduced
leaves, 7–10 mm long and 2–2.5 mm diam.; micro- No uses have been recorded of this rare species.
348 sporophylls triangular, ending in a lanceolate apex
ca. 1 mm long, with two basal pollen sacs partly hid-
den by the base of the microsporophyll. Seed cones Dacrydium gibbsiae Stapf, J. Linn. Soc., Bot.
mostly below end of foliage branchlets, terminal on 42: 192, t. 4. 1914. Type: Malaysia: Sabah, Ranau
very short dwarf shoots with reduced, 2–3 mm long, District, Mt. Kinabalu N. P., spur above Lobang,
narrowly triangular leaves; cones 4–5 mm long; cone L. S. Gibbs 4162 (holotype K). Fig. 115
bracts 3–4 mm long, becoming reddish. Seeds one
or two per cone; mature seeds not observed. Dacrydium beccarii Parl. var. kinabaluense Corner,
Gard. Bull. Straits Settlem. 10: 244, t. 9. 1939.
Taxonomic notes
Etymology
De Laubenfels (1988) described this as a new species
in his treatment of the conifers in Flora Malesiana, This species was named after Lilian Suzette Gibbs
distinguishing it from D. spathoides in having longer (1870–1925), who collected and studied the flora of
and straight leaves (as in juvenile plants) even on Mt. Kinabalu.
mature trees. Only the fertile structures (both male
and female) have adult type (much reduced) leaves
Vernacular names
at their base on very short dwarf shoots. The other
species in Borneo with juvenile type acicular leaves No vernacular names have been recorded.
has curved leaves which are not flat (not wider than
thick) terminating in an acute apex. This species is
Description
represented by a few specimens in the National Her-
barium Netherlands, Leiden branch (L), and one at Small trees 2–12 m tall, usually monopodial but with
Kew (K) and remains poorly known. low, spreading and assurgent primary branches; foli-
age branches in dense tufts towards ends of primary
branches, forming a flat-topped or domed candela-
Distribution
bra crown. Juvenile leaves only on seedlings and sap-
Borneo: Sarawak (Bukit Lawi, Merurong Plateau). lings, spreading widely or in young trees spreading
TDWG codes: 42 BOR-SR at 30° forward, covering shoot, 12–18 mm long, 1–1.2
mm wide, straight or slightly curved, keeled abaxi-
ally, pungent. Adult leaves robust, directed forward,
Ecology
densely covering shoot, linear to linear-lanceolate,
This species is locally common in primary mossy (3–)5–8(–10) mm long, 0.9–1.2 mm wide, sharply
forest on exposed mountain ridges between 1000 m keeled abaxially; apex incurved, pungent to apicu-
and 1500 m a.s.l. late. Stomata mostly on adaxial side in intermittent
lines from base to apex on both juvenile and adult
leaves, few near base on abaxial side. Pollen cones
Conservation
terminal or on a short lateral branchlet, cylindrical,
This species is only known from two localities, 20–25 × 4.5–7 mm; microsporophylls lanceolate, 4–5
where it is locally common. The type of forest it oc- mm long, 1.5 mm wide at base. Seed cones termi-
curs in and its habitat generally are not prime targets nal or often on short lateral branchlets with smaller
leaves; bract leaves longer, the upper one subtending Dacrydium gracile de Laub., Fl. Malesiana, ser. 1,
the seed nearly equally long as seed and spreading, 10 (3): 367. 1988, [“gracilis”]. Type: Malaysia: Sabah,
leaving the seed exposed. Seeds solitary or some- Ranau District, Mt. Kinabalu N. P., Silau-Silau trail,
times in pairs, obliquely positioned, ovoid, 4–5 mm D. J. de Laubenfels P 716 (holotype L). Fig. 116
long, covered for basal third by the epimatium, lus-
trous light brown.
Etymology
The species epithet refers to the thin, slender (juve-
Distribution
nile) leaves.
Borneo: Sabah (Mt. Kinabalu).
TDWG codes: 42 BOR-SB 349
Vernacular names
No common names have been recorded for this spe-
Ecology
cies.
This species occurs on Mt. Kinabalu from swampy
mossy forest at 1500–2000 m a.s.l. to high rocky
Description
ridges and slopes up to 3600 m. It is mostly restricted
to the serpentine and its ultramafic erosion products Trees to 40 m tall; trunk to 100 cm d.b.h., usually
where competition from other trees is limited. Be- with a clear bole and with long, tortuous branch-
low the summit of Mt. Kinabalu it grows as a shrub es, developing a broadly spreading and very open
on granite. The vegetation there is low and more or crown, sometimes more or less flat-topped. Bark
less open, with other conifers, e.g. Dacrycarpus im- scaly; outer bark dark brown; inner bark exuding
bricatus, Phyllocladus hypophyllus and Podocarpus red resin from slash. Foliage of two types, with ju-
gibbsiae, and Myrtaceae, e.g. Leptospermum recur- venile leaves and adult leaves markedly differing
vum, Tristaniopsis spp., Ericaceae and various small in length and width. Juvenile leaves remotely set,
oaks (Quercus) as codominants. At high altitudes the spreading at 50–90° from shoot, acicular, very slen-
common ericaceous shrub Rhododendron ericoides der, 12–20 × 0.3–0.4 mm, curved forward, triangular
has an uncanny resemblance to Dacrydium gibbsiae in cross section, pungent. Adult leaves more densely
until one sees the red tubular flowers. Codominant set, spreading at 30–45° forward, closer to shoot
or common conifers that grow with D. gibbsiae at on leading branches, acicular, slightly to strongly
higher altitudes on the mountain are Phyllocladus curved forward, (3–)4–6(–8) × 0.4–0.5(–0.7) mm,
hypophyllus, Dacrycarpus kinabaluensis, and Podo- triangular in cross-section and slightly wider than
carpus brevifolius. thick, weakly keeled abaxially, apiculate, dark green,
flushing light green. Stomata on all sides (leaves am-
phistomatic), in 1–3 intermittent lines on each face
Conservation
to apex. Pollen cones terminal or lateral on branch-
Although as far as is known this species is restricted lets with adult leaves, 6–7 × 2 mm at maturity; mi-
to Mt. Kinabalu, where it is abundant in specific hab- crosporophylls 0.7 mm wide at base, with extended
itats, no decline has been observed. Mount Kinabalu narrow apex 0.6–1 mm long, with two basal pol-
is a National Park which receives many visitors who len sacs, turning brown. Seed cones mostly below
hike up the mountain, and fire prevention might be- ends of foliage branches with adult leaves, terminal
come an issue if tourism increases. Little is known on dwarf shoots, subtended by shorter involucral
about the capacity of this species to regenerate after leaves, covered with bracts 2–3 mm long and reach-
fire. ing to base of seed, turning reddish at maturity.
IUCN: LC Seeds usually single, obliquely positioned, 3–3.5 mm
long, covered for the basal half by the epimatium
and turning lustrous dark brown to black.
Uses
No uses have been recorded of this species.
 Distribution Vernacular names
Borneo: Sabah (Mt. Kinabalu and vicinity), one lo- No vernacular name has been recorded for this spe-
cality in Sarawak. cies.
TDWG codes: 42 BOR-SB BOR-SR
Description
Ecology
Slender shrubs or dwarf trees to 2.5 m tall; stem
Dacrydium gracile is a tree occurring scattered in thickened at base, often sparsely branched. Bark
lower montane rainforest. It is usually associated rough, with horizontal stripes and large lenticels,
350 with the conifers Agathis borneeensis, Podocarpus breaking into small scaly plates, brown weather-
laubenfelsii, Sundacarpus amarus, Falcatifolium fal- ing blackish grey; inner bark fibrous, red-brown.
ciforme, Nageia wallichiana, and Dacrycarpus im- Branches spreading and ascending, with foliage
bricatus on soils poor in nutrients (kerangas forest); branches upright and tufted near ends. Leaves per-
in Sarawak it occurs in low canopy ‘heath’ forest on manently of the juvenile type, densely covering
sandstone. branches, acicular, curved forward or sometimes
curved near base only, (10–)13–15(–17) × 1 mm,
transverse triangular in cross-section, pungent.
Conservation
Stomata in 2–5 inter­mittent lines on all three sides
This species is quite rare and occurs as individual (higherst number of lines on adaxial side) from base
trees in primary forest. Most herbarium collections to apex. Pollen cones both terminal and lateral; ter-
are from Mt. Kinabalu and vicinity (partly in Mt. minal cones 8–15 mm long and 4 mm wide when
Kinabalu National Park) and the disjunct locality in expanded; lateral cones at base of a terminal cone
Sarawak (in a different habitat) could indicate that and smaller. Microsporophylls 4–5 mm long near
it is more widespread than currently known. With base of cone, with a greatly elongated acicular apex,
the present knowledge its area of occupancy (AOO) gradually shortening towards cone apex to 1–2 mm;
is certainly less than 20 km2 but the total number of pollen sacs 2, partly hidden by the curved basal part
mature individuals is unknown. of the microsporophyll. Seed cones terminal on long
IUCN: VU (D2) or short foliage shoots, 8–10 mm long, hidden by
leaves. Seeds 1–5 per cone, proximally covered for
a third to half by the epimatium and subtended by
Uses
bracts; bracts long, leaf-like, much exceeding the
No uses have been recorded of this species; it can be seed; seeds ovoid-oblong, 4–5 × 2.5 mm, brown,
assumed to be of value for its timber like other spe- with a constricted apex.
cies in the genus that grow into tall forest trees. Its
protected status within Mt. Kinabalu National Park
Distribution
make exploitation at least of these trees unlikely;
outside this protected area it may be logged. New Caledonia (Grand Lac, Lac en Huit, Rivière des
Lacs).
TDWG codes: 60 NWC
Dacrydium guillauminii J. T. Buchholz, Bull. Mus.
Hist. Nat. (Paris), sér. 2, 21: 282. 1949. Type: New
Ecology
Caledonia: Grande Terre, Province Sud, Chûtes de
la Madeleine, J. T. Buchholz 1728 (holotype ILL). Dacrydium guillauminii is a strictly riparian species
Fig. 117 growing on the banks and shores of streams and
small lakes that are frequently flooded; its seeds ger-
minate in water-logged soil. The Plaine des Lacs is
Etymology
a basin where small rivers and shallow lakes drain
This species is named after the French botanist An- ultramafic laterite soils and eroded serpentine hills,
dré Guillaumin (1885–1974). and the water after heavy rains is often of a milky
colour (one locality is on the aptly named Pernod
Creek). This species does not occur above the high
Vernacular names
water line. The altitudinal range is 150–275 m a.s.l.
It is commonly associated with Retrophyllum minus, No common names are known for this rare species.
another podocarp restricted to this habitat. Just me-
ters away where the ground rises, the ‘maquis mini-
Description
er’ vegetation dominates; consequently the niche of
this species is (literally) very narrow. Shrubs or trees [habit remained undescribed in
the protologue and subsequent treatments and has
not been observed by the author] with very slender
Conservation
ultimate foliage branchlets (0.25–0.4 mm diam.),
The extreme rarity of this species and its very re- including the leaves 1–2.5 mm diam. Leaves acicu- 351
stricted habitat alone put this species at risk of ex- lar, spreading from base but abruptly bent forward
tinction. The 2–3 populations are fragmented due below the middle parallel to the filiform branchlet
to the very local presence of its habitat; changes in or even curved inward, on ultimate branchlets 1–1.5
the water regime (e.g. by pollution or diversion of mm long, 0.2–0.4 mm wide, 0.1 mm thick, subulate,
streams caused by mining operations upstream) transverse-triangular, sharply keeled abaxially, flat
could destroy these. Although one population is or slightly concave on adaxial side, finely acumi-
protected in the Chûtes de la Madeleine Botanical nate; leaves on leading (older) branches adpressed,
Reserve, the others are unprotected and on land narrowly triangular, 2–3 mm long and 0.6–0.7 mm
under mining concessions (not actively mined at wide near base, dorsiventrally flattened, weakly
present). The species is very sensitive to fire and keeled on the abaxial side, acute-pungent. Fertile
numbers of individuals have declined, probably as material (pollen cones and seed cones) remaining
a result of increased tourism; restoration of habitats unobserved.
in the reserve is now being undertaken after tourism
became better regulated.
Taxonomic notes
IUCN: CR [B1ab(iii)+2ab(iii);C2a(i)]
This species is imperfectly known taxonomical-
ly as no fertile material has ever been collected.
Uses
Originally described as Podocarpus leptophyllus by
There are no recorded uses of this species and (ex- ­Wasscher (op. cit.) and transferred to Dacrycarpus
cept in a few botanical collections) it is not in culti- by Gaussen, it was finally settled in the genus Dacry-
vation. dium by De Laubenfels, where it undoubtedly be-
longs. The combination under Dacrydium was first
validly published by Silba as cited above. Wasscher
Dacrydium leptophyllum (Wasscher) de Laub. was uncertain of its placement for want of “flowers
ex Silba, Phytologia Mem. 7: 27. 1984. Podocarpus and fruits” and suggested it could belong in Dacry-
leptophyllus Wasscher, Blumea 4 (3): 414. 1941; dium, but he unfortunately chose the wrong genus
Dacrycarpus leptophyllus (Wasscher) Gaussen, (which was then treated in a much wider sense than
Trav. Lab. Forest. Toulouse T. 2, 1 (2, 20): 150 (nom. at present). This remains a poorly known species of
inval., Art. 33.4). 1974; Bracteocarpus leptophyllus which few if any specimens other than the type are
(Wasscher) A. V. Bobrov & Melikyan, Bjull. identified.
Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 59.
1998. Type: Indonesia: Papua, Pegunungan Maoke,
Distribution
Mt. Goliath, A. C. de Kock 39 (holotype BO).
New Guinea: Papua (Mt. Goliath).
TDWG codes: 43 NWG-IJ
Etymology
The species epithet (Greek lepto = slender, thin, nar-
Ecology
row) means ‘with slender leaves’.
Mossy heath forest.
 on all sides (leaves amphistomatic), but extending to
Conservation
distal part of leaf only on adaxial side, in a few inter-
Only known from one location on a mountain sum- mittent rows separated by the keel and by a midrib
mit. First described as Podocarpus leptophyllus by on the adaxial side. Pollen cones terminal or lateral
Wasscher (op. cit.), based on a herbarium collection below a terminal cone, 4–7 × 1.2 mm when mature;
made in 1911 and not recorded from other localities microsporophylls narrowly triangular, keeled abaxi-
since. Population size unknown, but likely fewer ally, tapering to an acute apex, with two basal pollen
than 1000 mature trees; no evidence of decline, but sacs. Seed cones terminal, often on dwarfed shoots,
obviously at some risk if this locality became subject 2–3 mm long, subtended by short (1–2 mm) leaves
to environmental changes detrimental to this spe- followed by longer (2–3 mm) flattened cone bracts
352 cies. surrounding the basal epimatium, becoming swol-
IUCN: VU (D1, 2) len and red at base. Seeds single, covered for only
the basal third or less by the epimatium, 3.5–4 mm
long, ovoid but slightly flattened, lustrous brown.
Uses
There are no uses recorded of this species.
Distribution
New Caledonia: Grande Terre, Province Sud.
Dacrydium lycopodioides Brongn. & Gris, TDWG codes: 60 NWC
Bull. Soc. Bot. France 16: 329. 1869. Type: New
Caledonia: Grande Terre, Province Sud, Mt. Mou,
Ecology
J. A. I. Pancher s.n. (holotype P).
Dacrydium lycopodioides is an uncommon tree in
montane forest on serpentine, occurring in a few
Etymology
sheltered ravine forests or hollows which have es-
The species epithet refers to similarity of the foli- caped the ravages of fire in the surrounding land-
age branches with Lycopodium, a genus of clubmoss scape. These forest pockets are naturally less prone to
common in New Caledonia. burn than the drier woodland and ‘maquis minier’,
but are not absolutely safe from forest fires. This spe-
cies also occurs on moist mountain slopes and ridg-
Vernacular names
es (e.g. Mt. Mou) where fires are infrequent. Here its
No vernacular name is known for this species. stature becomes more stunted to 5–10 m as it occurs
towards the summit area at around 1000–1200 m
a.s.l. in association with the conifers Araucaria hum-
Description
boldtensis, A. laubenfelsii (emergents), Falcatifolium
Trees to 25 m tall; trunk d.b.h. to 50 cm, erect, in for- taxoides, and Prumnopitys ferruginoides in low for-
est forming a clear bole; crown spreading, dense and est with numerous angiosperms among which spe-
rounded in old trees. Bark scaly with small flakes, cies of the Myrtaceae are dominant.
brown, weathering grey; inner bark more or less fi-
brous and light brown. Foliage in much branched
Conservation
tufts, spreading to erect in mature trees. Leaves
on seedlings and juvenile trees acicular, 7–10 mm This species is known from several herbarium col-
long, 0.2–0.3 mm wide and thick, spreading at ca. lections made over a period of more than 150 years,
45° from shoot, slightly curved forward to nearly some of which are without locality data. The locali-
straight, triangular in cross section, pungent. Adult ties recorded are Mt. Mou, Mt. Dzumac, Mt. Naka-
leaves shorter and gradually changing from juvenile da, Mt. Humboldt, and a mountain ridge above the
leaves, spirally arranged and spreading at 30–40°, Ouinné River; it was most frequently collected from
curved forward above base, linear-lanceolate, (2–) Mt. Mou. This may partly be due to relative ease of
3–5(–6) mm long, 0.6–0.8 mm wide just above base access of the latter mountain and it may occur else-
and 0.3–0.4 mm thick, keeled sharply on abaxial where in the mountains of the SE part of Grande
side, gradually tapering to a pungent apex. Stomata Terre. It is obviously a relatively rare species, but it is
not at risk at present, thanks to the relatively undis- and shape to adult leaves; the cone bracts longer, 6–8
turbed mountain ravines in which it occurs. mm, spreading or curved inward partly covering the
IUCN: NT seed. Seeds solitary or in pairs, obliquely positioned,
covered for basal third by the epimatium, 4–5 mm
long, ovoid, brown.
Uses
No commercial or other uses are recorded of this
Taxonomic notes
rare species.
This species is quite similar in its botanical char-
acters to D. medium, described by the same author
Dacrydium magnum de Laub., J. Arnold Arbor. 50: from Malaya. Rather subtle differences in the adult 353
299. 1969. Type: Indonesia: Maluku, Obi, G. A. L. de foliage leaves lead however to a different appearance
Haan bb 23.806 (holotype L). of branchlets; the pollen cones of D. magnum are
longer and the two species occupy different habi-
Dacrydium beccarii Parl. var. rudens de Laub., J. Ar- tats, leading to greater tree size in D. magnum. The
nold Arbor. 50: 303. 1969. fact that the distributions of the two species do not
overlap is in the Malesian Archipelago of less sig-
nificance; disjunct distribution in this genus within
Etymology
species and even varieties is rather commonplace, as
The species epithet means ‘large’ and was presum- it is among other conifers.
ably given for its relatively large pollen cones.
Distribution
Vernacular names
Malesia: Maluku [Moluccas] (Obi Island); Papuasia:
No common names are recorded for this species. Louisiades Archipelago (Tagula Island), Solomon
Islands.
TDWG codes: 42 MOL 43 NWG-PN SOL-SO
Description
Trees to 30 m tall; trunk d.b.h. to 60 cm, monopodial,
Ecology
crown compact, densely branched. Bark scaly, red-
dish brown; inner bark pinkish or reddish. Prima- Dacrydium magnum is a canopy tree in tropi-
ry branches spreading and ascending, terminating cal rainforest at altitudes between 60 m and 1200
with dense tufts of upright foliage branches. Juvenile m a.s.l.; on ridges of mountains it can become the
leaves spreading widely, similar to adult leaves but dominant forest tree. On exposed sites it becomes
longer, to 20 mm, acicular, slightly curved, trian- more stunted, but not a low shrub.
gular in cross section, keeled abaxially, 0.4–0.5 mm
wide; apex pungent. Adult leaves slightly spread-
Conservation
ing but strongly directed forward on shoot and
more or less imbricate, short acicular, (2–)3–5(–6) The conservation status of this species is difficult to
× 0.3–0.4 mm, mostly uniform on a shoot, curved assess due to its wide and scattered distribution. It
with apex turning back to the shoot (making it pos- is known from islands west and east of New Guinea
sible to strike a finger tip backwards over a foliage but not from New Guinea itself. It is undoubtedly a
branchlet), triangular in cross section, keeled abaxi- rare species which may well turn up in collections
ally, apiculate. Stomata on all sides (leaves amphi­ made in New Guinea already or to be made in fu-
stomatic) but more numerous on the adaxial side, ture. Due to the possibly wider distribution than
in intermittent lines to apex. Pollen cones terminal presently known, but at the same time the likely-
on foliage branchlets, 10–16 × 2–2.5 mm. Micro­ hood of exploitation along with other species of Po-
sporophylls with elongated and strongly incurved docarpaceae from which loggers will not distinguish
apex and two basal pollen sacs. Seed cones terminal, it, it is here listed as Near Threatened.
ca. 3 mm long, subtended by leaves similar in length IUCN: NT
 Uses Distribution
No uses have been recorded for this species. Malesia: Peninsular Malaysia, N Sumatera.
TDWG codes: 42 MLY-PM SUM

Dacrydium medium de Laub., Blumea 23 (1): 98.

Ecology
1976. Type: Malaysia: Peninsular Malaysia, Pahang,
Gunung Tahan, D. J. de Laubenfels P 540 (holotype Dacrydium medium occurs mostly on mountain
L). summits and ridges in low, scrubby vegetation on
poor soils that will not support forests. The soils in
354 Dacrydium beccarii Parl. var. subelatum Corner, these places are often very shallow and rocky. The
Gard. Bull. Straits Settlem. 10: 243, t. 7. 1939. altitudes of these sites vary between 1350 m and 2100
m a.s.l. The conifers are either shrubs themselves or
small trees rising above the general vegetation, in
Etymology
which Myrtaceae (e.g. Baeckea and Leptospermum)
The species epithet describes it as being intermedi- are most common. More rarely scattered individu-
ate (midway between) two other species. als occur in forest below the mountain ridges where
they may grow taller to keep their crowns in or
above the canopy as they are light demanding.
Vernacular names
sangu (Gajo Lands, Sumatera)
Conservation
IUCN: LC
Description
Shrubs or small, often gnarled trees to 4–6 m tall,
Uses
some trees in forest to 20 m, crown compact, dense-
ly branched. Bark scaly, brown. Primary branches No uses are recorded of this species.
spreading and ascending, terminating with dense
tufts of upright foliage branches. Juvenile leaves
spreading, acicular, similar to adult leaves but long- Dacrydium nausoriense de Laub., J. Arnold Arbor.
er, 10–20 mm, slightly curved, transverse-triangular 50: 287. 1969, [“nausoriensis”]. Type: Fiji: Viti Levu,
in cross section, keeled abaxially, 0.5–0.6 mm wide; Navosa, Nausori Highlands, 6.5 km SE of Nausori,
apex pungent. Adult leaves spreading slightly, di- D. J. de Laubenfels P 302 (holotype A).
rected forward on shoot and more or less imbricate
on leading shoots, acicular, (2–)4–6(–10) × 0.5–0.6
Etymology
mm, mostly uniform on a shoot, nearly straight with
incurved apex, on older plants up to 6 mm long, The species epithet refers to the Nausori Highlands
more strongly curved inward, triangular in cross on the island of Viti Levu from where this species
section, keeled abaxially, apiculate. Stomata on all was described.
sides (leaves amphistomatic) but more numerous
on adaxial side, in intermittent lines to apex. Pol-
Vernacular names
len cones terminal on foliage branchlets, 7–9 × 2.5
mm; microsporophylls with elongated and strong- yaka, tangitangi (Fijian)
ly incurved apex and two basal pollen sacs. Seed
cones mostly terminal, ca. 3 mm long, subtended by
Description
shorter leaves to 2 mm long; cone bracts longer, 3–5
mm long, spreading or curved inward partly cover- Trees of medium size, to 25 m tall; trunk to 50 cm
ing the seed, flattened and turning reddish at matu- d.b.h., much branched eventually developing a
rity. Seeds solitary, obliquely positioned, covered for broadly spreading domed crown. Bark scaly with
basal third by the epimatium, 4–5 mm long, ovoid, large flakes; outer bark brown weathering grey; in-
brown to black. ner bark pink to reddish. Foliage of two types, with
juvenile leaves and with adult leaves. Juvenile leaves former rainforest which was the habitat of this spe-
acicular, on seedlings and saplings very slender, 10– cies has been converted to agriculture or pasture,
15 × 0.3 mm, shorter and slightly thicker on young especially in more densely settled areas. Regenera-
trees, straight or slightly curved forward, spreading tion is slow and does not appear to be adequate; this
at more than 45° from shoot, keeled on four sides, podo­carp probably needs periodic disturbance by
pungent. Transitional leaves present on saplings, 3–5 fire to spread from seed, but too frequent fires de-
mm long, spreading at ca. 45°. Adult leaves on nu- stroy that opportunity.
merous slender, much branching shoots, decurrent IUCN: EN (A1cd, B1e)
at base, spreading out and forward at ca. 30° from
shoot, short acicular, 1–1.5 mm long and 0.3–0.4 mm
Uses
wide (on leading branches slightly larger), keeled 355
abaxially, obtuse or sometimes acute with incurved Considered a highly valuable timber tree in Fiji
apex. Stomata on all sides (leaves amphistomatic), (graded Class I), this species is used in construction
on juvenile leaves in narrow grooves, on adult leaves and all kinds of carpentry work.
in a few scattered lines, absent on the distal part of
the abaxial side, on the adaxial side to apex. Pollen
cones terminal or lateral on branchlets with adult Dacrydium nidulum de Laub., J. Arnold Arbor. 50:
leaves, 4–6 × 1–1.2 mm at maturity, microsporo- 292. 1969. Type: Indonesia: Papua, Jazirah Doberai,
phylls triangular, 1 × 0.7 mm, acute-apiculate, with Lake Aja-Maroe, W side of lake [“Segior”], W. Vink
two basal pollen sacs. Seed cones terminal on long & M. Vink BW 15271 (holotype L).
or very short branchlets with adult leaves, ca. 3 mm
long, with distal bracts 2 mm long, slightly swelling
Etymology
and turning red at maturity. Seeds solitary, free from
leaves, obliquely positioned, covered for basal third The species epithet is derived from Latin nidus (=
by the epimatium, ovoid, 3.5–4 mm long, turning nest) and refers to the ‘nesting’ of the growing seed
lustrous brown. cone within its elongated leaf-like bracts.

Distribution Vernacular names

Fiji (Vanua Levu, Viti Levu). yaka (Fijian)
TDWG codes: 60 FIJ
Description
Ecology
Trees to 30 m tall, monopodial and erect; trunk
Dacrydium nausoriense is a canopy tree in more d.b.h. to 70 cm or rarely more, usually with a clear
or less open montane rainforest (annual rainfall bole; crown spreading, dense and rounded in old
ca. 3000 mm) and fairly common on the plateaux trees. Bark irregularly fissured, scaly, dark brown,
and summits of the Nausori Highlands. It is asso- weathering grey; inner bark slightly fibrous and red-
ciated with Dacrycarpus imbricatus and Podocarpus brown. Foliage in much branched tufts, drooping to
degeneri and with numerous angiosperm trees and spreading in mature trees. Leaves on seedlings and
shrubs. juvenile trees long acicular, 10–20 mm long, 0.2–
0.3 mm wide and thick, spreading widely, slightly
curved to nearly straight, triangular in cross section,
Conservation
pungent. Adult leaves much shorter but variable in
This species exists in small stands in remnants of length between trees and within trees, spirally ar-
rainforest and is exploited for its timber. Much of ranged and decurrent at base, short acicular, (1.5–)
this forest is privately owned or on tribal land or 3–5(–7) mm long, 0.3–0.7 mm wide and 0.3–0.4 mm
both and there are no forest reserves where this spe- thick, keeled sharply on abaxial side, spreading at
cies is being protected. It is mostly known from the 30–45°, slightly curved or nearly straight but with an
Nausori Highlands but other localities have been incurved, obtuse to short apiculate apex. Intermedi-
found, one of them on Vanua Levu. Much of the ate forms of leaves occur on younger trees. Stomata
 on all sides (leaves amphistomatic), but extending to Dacrydium novoguineense Gibbs, Contr.
apex only on the adaxial side, in several intermittent Phytogeogr. & Fl. Arfak Mts.: 78. 1917. Type:
rows. Pollen cones terminal or lateral, ca. 4 × 2 mm Indonesia: Papua, Arfak Mts., Koebre Ridge,
when immature, elongating to 18 mm; microspo- L. S. Gibbs 5648 (lectotype BM).
rophylls triangular, 1.2 × 0.8 mm, keeled abaxially,
apiculate, with two basal pollen sacs. Seed cones ter-
Etymology
minal, 2–3 mm long, subtended by short (1.5–2 mm)
leaves, followed by longer (3–5 mm) cone bracts The species epithet refers to its occurrence in New
surrounding the epimatium, becoming swollen and Guinea.
red at base. Seeds solitary, covered for only the basal
356 third or less by the epimatium, 3.5–4 mm long, lus-
Vernacular names
trous brown.
kaowié, kowié, aru, munump (New Guinea, various
languages)
Taxonomic notes
This species is very similar to Dacrydium lycopodio-
Description
ides of New Caledonia, but differes in having much
longer pollen cones and non-flattened bracts, which Trees of small to large size, to 30 m tall; trunk to
encase the growing seed cone as if in a nest. 50 cm d.b.h., much branched eventually developing
a broadly spreading domed crown, sometimes more
or less flat-topped. Bark scaly; outer bark brown
Distribution
to blackish; inner bark reddish, slightly resinous.
Malesia: Maluku [Moluccas] (Halmaheira), Su- Foliage of two types, with juvenile leaves and with
lawesi, Lesser Sunda Islands (Sumba); Papuasia: adult leaves. Juvenile leaves acicular, on seedlings
D’Entrecasteaux Islands, Louisiades Archipelago, and saplings 5–12 × 0.4–0.6 mm, shorter on young
New Guinea; SW Pacific: Fiji. trees (especially at base of branchlets), curved for-
TDWG codes: 42 LSI-LS MOL SUL 43 NWG-IJ NWG- ward, spreading at less than 45° from the shoot,
PN 60 FIJ strongly keeled abaxially, pungent. Transitional
leaves sometimes present on mature trees, but usu-
ally abruptly changing to adult leaves, also present
Ecology
on saplings less than 1 m tall. Adult leaves scale-like
Dacrydium nidulum is a canopy tree of mostly low- on slender, cord-like, much branching shoots 2–7
land tropical rainforest, from 10 m to 750 m above cm long and 0.8–2.3 mm wide (ultimate branchlets
sea level and rarely to 1200 m. It often occurs along variable in width, thread-like or more robust), ap-
rivers or on lake shores and regenerates well in sec- pressed, oblong-rhombic, 1.3–2 × 0.3–0.5 mm on ul-
ondary forest after disturbances. It is found on soils timate branchlets, slightly larger on leading shoots,
over limestone as well as acidic types of rock. rounded or keeled abaxially, obtuse or acute with
incurved apex. Stomata on all sides (leaves amphi­
stomatic), on juvenile leaves in narrow grooves, on
Conservation
adult leaves in scattered lines, mainly on adaxial
IUCN: LC side. Pollen cones terminal on branchlets with adult
leaves (sometimes with transitional leaves), 5–8 × 1.5
mm at maturity; microsporophylls triangular, 1.2 ×
Uses
1 mm, obtuse or apiculate, with two basal reddish
In Fiji, where this species (and D. nausoriense, not pollen sacs, turning brown. Seed cones terminal on
distinguished by loggers) is considered a very valu- branchlets with adult leaves, 3 mm long, covered
able timber tree, it is used mainly for construction with elongated bracts to 3 mm long, turning red at
and carpentry such as furniture making. maturity. Seeds solitary, obliquely positioned, cov-
ered only at base by the epimatium, 4–5 mm long,
turning dark brown.
 spreading, dense and rounded in old trees. Bark
Distribution
brown, weathering grey, peeling profusely in broad,
New Guinea. loose brown strips. Foliage in much branched tufts,
TDWG codes: 43 NWG-IJ NWG-PN drooping to more or less erect on mature trees.
Leaves on seedlings and juvenile trees long acicular,
10–20 mm long, 0.2–0.4 mm wide and thick, slight-
Ecology
ly curved, abaxially keeled, pungent. Adult leaves
Dacrydium novoguineense is very common in New much shorter but variable in length between trees
Guinea, where it forms a canopy tree in open forest and within trees, linear, (1.5–)3–5(–7) mm long, 0.4–
especially on mountain ridges. In forests at middle 0.8 mm wide and thick, keeled sharply on abaxial
altitudes (1500–2200 m) it is an emergent above the side, spirally arranged and decurrent at base where 357
general canopy, but higher up on ridges it becomes they are imbricate, spreading at 30–45°, slightly
a small tree among other scrub and tree ferns. It of- curved or nearly straight but with an incurved, ob-
ten regenerates abundantly after disturbance by fire, tuse to short apiculate apex. Intermediate forms of
when it can become temporarily dominant. This leaves present on younger trees. Stomata on all sides
species is indifferent to soil types and occurs on clay, (leaves amphistomatic), but extending to apex only
sand, quartzite and other rock debris as well as on on adaxial side, in several intermittent rows. Pollen
peat. cones terminal, ca. 4 × 2 mm when immature, elon-
gating to 12 mm; microsporophylls narrowly trian-
Conservation gular, 1.5 × 0.5 mm, keeled abaxially, apiculate, with
two basal pollen sacs. Seed cones terminal, subtend-
IUCN: LC
ed by short leaves followed by slightly longer (2–3
mm) cone bracts, becoming swollen and red. Seeds
Uses
single, covered for only the basal third or less by the
The timber of this species is used for building and epimatium, ovoid, 4–5 × 3 mm, chestnut to lustrous
the bark for insulation of the walls of traditional dark brown.
houses in the highlands of New Guinea.
Taxonomic notes
Dacrydium pectinatum de Laub., J. Arnold Arbor. This species resembles the lowland form of D. nid-
50: 289. 1969. Type: Malaysia: Sabah, Beaufort ulum, but differs from it by its more robust leaves
District, Sandakan, D. I. Nicholson SAN 17292 and by the fully exposed mature seed. The length of
(holotype A). leaves in mature trees is influenced by environmen-
tal conditions, with shorter leaves associated with
Dacrydium pectinatum de Laub. var. robustum de more adverse environments such as peat bogs.
Laub., J. Arnold Arbor. 50: 291. 1969.
Distribution
Etymology
China: Hainan; Malesia: Borneo (including Karima-
The species epithet refers to the spreading (pecti- ta & Natuna Islands), Philippines, Pulau Belitung.
nate) arrangement of the foliage leaves. TDWG codes: 36 CHH 42 BOR-BR BOR-KA BOR-SB
BOR-SR PHI SUM
Vernacular names
Ecology
malur, melur, sempilor, tjemantan (different parts of
Borneo); lokinay (Philippines). Dacrydium pectinatum is a species of lowland to
montane rainforests, where it occurs as scattered
individual trees at altitudes from near sea level to
Description
1500 m a.s.l.; above 600–800 m trees become scarce.
Small to medium size trees to 20 m, rarely to 40 More dense stands or even pure stands of this spe-
m tall; trunk d.b.h. to 50 cm or rarely more; crown cies are restricted to nutrient-poor soils like shallow,
 leached sands or to swamps with peat formation both sides (leaves amphistomatic) but more abun-
above the water table. The forest on nutrient-poor dant on the adaxial side where lines extend to apex.
sandy soils are known as kerangas and the poor- Pollen cones not observed. Seed cones terminal, of-
est sites may only support ‘heath forest’ dominated ten on a small lateral shoot, 2–3 mm long, subtended
by this conifer and Gymnostoma (Casuarinaceae) by reduced leaves less than 2 mm long; cone bracts
which looks in habit like a conifer. In Sabah D. pec- slightly spreading, straight, up to 3 mm long and 0.5
tinatum may occur on ultrabasic soils supporting mm wide, covering the basal epimatium of the seed
an open, low vegetation of shrubs and ferns, where when mature. Seeds solitary or in pairs, at oblique
trees remain small. angle to cone or shoot apex, ovoid, slightly flattened,
ca. 4 mm long, brown.
358
Conservation
Taxonomic notes
IUCN: LC
De Laubenfels (1969), when first describing this new
species, included collections from Borneo (Sarawak)
Uses
in it, but later, in Flora Malesiana (De Laubenfels,
No uses have been recorded for this species. 1988) he had concluded that these belong to Dacry-
dium ericoides (E. F. Brunig SFN 8722, type). The
latter species has longer and slightly wider, straight
Dacrydium spathoides de Laub., J. Arnold Arbor. and distinctly flattened leaves narrowing abruptly to
50: 299. 1969. Type: Indonesia: Papua, Pegunungan an apiculate apex. Hence D. spathoides, as far as is
Maoke, Idenburg River, 18 km SW of Bernhard known, is restricted to one locality in New Guinea.
Camp, L. J. Brass 12659 (holotype A).
Distribution
Etymology
New Guinea: Papua (Central Highlands, Idenburg
The species epithet refers to the shape of the adult River drainage).
leaf resembling a spathe (as in palms) with its con- TDWG codes: 43 NWG-IJ
cave adaxial side.
Ecology
Vernacular names
Dacrydium spathoides is known from a single local-
No common names have been recorded for this spe- ity on the Idenberg River where it forms a canopy
cies. tree in moist, mossy montane rain forest at an alti-
tude of 2150–2200 m a.s.l.
Description
Conservation
Trees 25–35 m tall; trunk monopodial, erect, to 50
cm d.b.h. Bark scaly, brown weathering grey; inner This species is known from only one locality and its
bark reddish and exuding red sap. Crown spread- type specimen, but due to poor collecting records
ing above a clear bole in canopy forest. Juvenile and from the area nothing can be concluded from this
adult leaves similar, mainly differing in size and with as to rarity, population size etc. The fact that is has
a gradual transition. Juvenile leaves flattened acicu- been described in 1969 and no subsequent collec-
lar, spreading at ca. 60°, 5–7 mm long, 1 mm wide tions were made is due to its obscurity as a species,
and 0.2–0.3 mm thick, straight or slightly curved which is not commonly known to botanists who
forward at the pungent apex, keeled on abaxial might visit the area. Rather than maintain it as DD,
side, slightly concave on adaxial side. Adult leaves the Conifer Specialist Group believes this species
linear-lanceolate, spreading at ca. 45°, 2–4 mm long, should be flagged as NT as it is surely very rare and
0.8–0.9 mm wide and 0.2–0.3 mm thick, straight restricted. As a canopy tree in rain forest it may be
or slightly curved forward at apiculate apex, keeled subject to logging.
abaxially, distinctly concave adaxially. Stomata on IUCN: NT
 Uses Taxonomic notes
The timber of this tree is possibly used for construc- Plants with foliage characters intermediate between
tion (house building), but its very localized occur- those of Dacrydium araucarioides and D. guillaumi-
rence means that no specific data are available. nii were first noted in 1994 by Bernard Suprin in the
area of the Chûtes de la Madeleine along the Riv-
ière des Lacs. These plants were observed there in
Dacrydium ×suprinii Nimsch, Feddes Repert. subsequent years, but rarely collected. A study by
118 (1–2): 52. 2007. Type: New Caledonia: Grande Knopf et al. (2007) investigating both macro- and
Terre, Province Sud, Chûtes de la Madeleine, 100 micromorphology concluded that these plants are
m downstream from waterfall, H. Nimsch 5122 a fixed hybrid species originating from these two 359
(holotype B). species, with D. guillauminii as the putative female
parent. Investigations into the DNA of all three taxa
would be required to test the hybrid hypothesis, but
Etymology
the morphological evidence, together with its close
This hybrid species was named after Bernard Suprin, proximity to the maternal parent, are strongly sup-
a French resident botanist with intimate knowledge portive of it.
of the New Caledonian flora.
Distribution
Vernacular names New Caledonia (Grand Lac, Rivière des Lacs).
TDWG codes: 60 NWC
No common names are known for this species.
Ecology
Description
Dacrydium ×suprinii is a semi-riparian species
Small shrub-like tree 2–4 m tall. Branches in mature growing on the banks and shores of streams and
plants in pseudo-whorls of a few first order branches small lakes that are frequently flooded; however,
only, spreading and ascending to form a candelabra unlike one of its parents, D. guillauminii, it may be
crown with tufts of foliage branches. Leaves have found on slightly drier ground in the transition to
characters intermediate between D. araucarioides low gallery forest along the river. It is commonly as-
and D. guillauminii; juvenile leaves 10–15 mm long, sociated with D. guillauminii, while the other parent,
0.8–1 mm wide, adult leaves densely arranged, 5–10 D. araucarioides, occupies higher and drier ground
mm long, ca. 1 mm wide, rhombic in cross-section, in ‘maquis minier’ and low gallery forest, a small
partly imbricate but free at least towards apex, nar- distance away from the water line.
rowly lanceolate with a wide base, gradually taper-
ing towards the acute-acuminate but non-pungent
Conservation
apex. Stomata few near base on abaxial (lower) face
and in two lateral lines from base to apex on the This hybrid taxon is very rare and so far only known
adaxial side. Pollen cones both terminal and lateral, from three localities in the drainage of the Rivière
10–16 mm long and 4–5 mm wide when expanded; des Lacs, within 15 km from each other. Its distribu-
lateral cones at base of a terminal cone and smaller. tion coincides closely with one of the parent species,
Microsporophylls with a greatly elongated acicu- D. guillauminii, but far fewer individuals appear
lar apex 4–5 mm long near base of cone, gradually to exist of the hybrid. The female parent species
shortening towards cone apex to 1–2 mm; pollen (D. guillauminii) is considered Critically Endan-
sacs 2, partly hidden by curved basal part of micro- gered (CR) on the IUCN Red List. At present, only
sporophyll. Seed cones terminal on long or short fo- one of the three localities is within a nature reserve:
liage shoots, ca. 6 mm long, hidden by leaves. Seeds the Chûtes de la Madeleine Botanical Reserve. This
1–2 per cone, proximally covered for half or more by reserve is surrounded by land under mining conces-
the epimatium and subtended by bracts; bracts leaf- sions for nickel; however, no operations are carried
like, long, exceeding the seed; seeds ovoid-oblong, out at the present time.
3–4 × 2 mm, with a constricted apex, brown. IUCN: CR (D)
 normal size leaves, with several spreading green
Uses
bracts. Seeds 1(–2), exposed, covered at the base by
No uses are recorded of this species and it is not the epimatium, ovoid, 4–5 mm long, lustrous brown.
known to be in cultivation.
Taxonomic notes
Dacrydium xanthandrum Pilg., Bot. Jahrb. Syst. In Flora Malesiana ser. 1, 10 (3): 369–370 (De Lauben-
69: 252. 1938. Type: Papua New Guinea: Morobe, fels, 1988) this species has been stated to occur in
Ogeramnang, M. S. Clemens 4504 (lectotype A). Peninsular Malaysia (“Malaya”) but critical exami-
Fig. 118, 119 nation of all herbarium specimens involved (FRIM,
360 K, L) has revealed that this was based on erroneous
identification: all specimens from the peninsula be-
Etymology
long to D. beccarii.
The species epithet is a composite word with Greek
xantho- = yellow and -andrus = male; it describes
Distribution
the colour of the pollen cones.
Malesia: Borneo, Maluku [Moluccas], Philippines,
Sulawesi, N Sumatera; Papuasia: New Britain, New
Vernacular names
Guinea, Solomon Islands (Bougainville).
kerapui (Sabah); arun gunong (Sumatera, Aceh); TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
hinubayan (Philippines) MLY-PM MOL PHI SUL SUM 43 BIS NWG-IJ NWG-PN
SOL-NO
Description
Ecology
Trees to 35 m tall, sometimes shrubs; trunk of trees
monopodial, to 80 cm d.b.h. Bark scaly and flaking; Dacrydium xanthandrum is a widespread and lo-
outer bark dark brown; inner bark red-brown exud- cally common or dominant species, occurring from
ing red resin. Branches spreading or assurgent and primary lowland rainforest with scattered individu-
foliage branches numerous, spreading or drooping, al tall trees to high montane ridges (up to 2700 m)
forming a dense, rounded crown. Juvenile leaves with low mossy forest, where it may dominate. Soils
and adult leaves very similar; juvenile leaves spread- vary from clay to peat but D. xanthandrum becomes
ing perpendicular to branch or curved forward from abundant or dominant only in exposed rocky ridges
base, to 20 mm long and 0.8–1 mm wide, strongly or in peat over acid rocks (dacite, granite, or sand-
keeled abaxially, ca. 0.2 mm thick without keel, nar- stone) or nutrient-poor sand. In such localities it is
rowly acute. Adult leaves short decurrent, crowded accompanied by various species of Myrtaceae and
but spreading perpendicular to shoot or at least at a other angiosperm shrubs and small trees.
wide angle, curving forward or nearly straight, (5–)
7–10(–13) mm long, 0.5–0.8 mm wide, slightly flat-
Conservation
tened with a sharp abaxial keel and a narrow adaxial
midrib, distally tapering to a pungent apex. Stomata IUCN: LC
on both sides (leaves amphistomatic), but in several
lines on either side of midrib on adaxial side and
Uses
only few on abaxial side near base. Pollen cones lat-
eral or terminal on foliage branches, subtended by The timber of this species is not distinguished from
a few short leaves ca. 2 mm long, 6–13 × 2–2.5 mm; other species of Dacrydium and where large trees
microsporophylls extended into a curved, rostrate occur in the forest, they will probably be logged and
apex, 1.5–2 mm long, yellow with two reddish basal traded as ‘sempilor’ a collective name for several
pollen sacs. Seed cones few, often below ends of foli- genera and species of Podocarpaceae. This wood is
age branches, terminal on a dwarf shoot with small, valued for construction and furniture making.
2–3 mm long leaves, or when terminal ­following
Diselma Hook. f., Fl. Tasmania 1 (5): 353. 1857. Type: Diselma archeri Hook. f.
(Cupressaceae).

Microcachrys W. Archer, Hooker’s J. Bot. 2: 51. 1850, and obtuse apex, 1–2 × 0.6–1.5 mm; stomata in two
non Hook. f. (1845). Type not designated. short whitish bands in lower part of the leaf, eglan-
dular; young leaves on mature branchlets reddish at
Greek: di = two; selma = upper deck, seat; referring first, turning green or olive-green. Pollen cones nu-
to the fertile scales in the seed cone. merous, terminal and solitary on ultimate branch-
lets, to 3 mm long; microsporophylls 6–8, smaller
than leaves, triangular-deltoid, usually 4 fertile, each 361
Description
with 2 abaxial pollen sacs. Seed cones terminal on
See the species description. ultimate branchlets, maturing in one season to small
cones 3 × 2 mm, consisting of 2 pairs of decussate
cone scales surrounding a central columella; upper
Distribution
scales fertile, 2–3 × 1–1.5 mm, with bract margin visi-
As for the species. ble at the distal end; columella relatively large, resin-
ous glandular, clavate or ovate-cylindrical, breaking
off in old persistent cones. Seeds 2 (or 1) developing
Diselma archeri Hook. f., Fl. Tasmania 1 (5): 353, t. on each fertile bract-scale complex, ovoid-flattened,
98. 1857. Type: Australia: Tasmania, Great Western brown; wings 2(–3), marginal, broad, partly sur-
Tiers, Meander Falls, [“summit of the western rounding seed.
mountains and Falls of Meander”], R. C. Gunn
[366] s.n. (lectotype K). Fig. 120
Taxonomic notes
The foliage and leaf arrangement as well as leaf mor-
Etymology
phology of Diselma archeri very closely resemble
This species was named after William Archer (1820– those of the sympatric prostrate shrub Microcachrys
1874), who studied the flora of Tasmania. tetragona (Podocarpaceae), which are opposite-
decussate (“in distichous pairs” according to Hill &
Brodribb, 1999), unlike the usually helical arrange-
Vernacular names
ment in that family. It may be worthwhile to investi-
Cheshunt pine gate whether the two taxa really have a homologous
phyllotaxis or whether the podocarpaceous conifer
has in fact a bijugate phyllotaxis (Tomlinson & Za-
Description
charias, 2001) as a deviation from a helical arrange-
Shrubs, or very small trees 1.5–4 m, occasionally ment. It is interesting that the terminal cones of both
a small tree up to 6 m in shade, rarely prostrate, sexes in Microcachrys have their sporophylls in a
stem to 30–40 cm diam. but mostly smaller, ever- helical arrangement, while those in Diselma are, like
green, dioecious; trunk monopodial, short, or mul- their leaf homologues, decussately arranged.
tistemmed, with sprawling or assurgent branches
from a common base, forming extensive bushes.
Distribution
Bark scaly, rough, brown turning grey; inner bark
reddish brown. Branches short, rigid, forming a W Tasmania, in most highland areas but absent in
dense crown. Foliage branches numerous, not in a the NW.
plane, very dense, straight and more or less rigid, TDWG codes: 50 TAS
short (10–20 mm), tetragonal or occasionally trigo-
nal, ca. 1.5 mm diam., covered with closely appressed
Ecology
leaves, persistent. Leaves opposite-decussate, occa-
sionally in whorls of 3, imbricate, scale-like, rhom- This shrubby species occurs in subalpine and alpine
bic, more or less keeled distally, with entire margins moorland and boulderfields, sometimes ­riparian
 along streams or on shores of lakes and ponds, almost all populations are reserved within the West-
among perennial herbs and other shrubs (‘conif- ern Tasmania Wilderness World Heritage Area
erous heath’), e.g. Astelia alpina, Pherosphaera ­(WTWWHA). It is not considered threatened, al-
hookeriana, Microcachrys tetragona, Podocarpus though it is sensitive to fires, which have in historic
lawrencei, Richea scoparia, R. pandanifolia, Loma- times increased in frequency in some areas.
tia spp., Orites acicularis, O. revoluta, Helichrysum IUCN: LC
spp., Leptospermum spp. and larger shrubs or stunt-
ed trees, e.g. Athrotaxis cupressoides, Phyllocladus
Uses
­aspleniifolius, Nothofagus gunnii, and Eucalyptus
coccifera. The altitudinal range of Diselma archeri is This shrub is uncommon in cultivation, but some
362 from 550 m to 1400 m a.s.l. Soils are acidic, often specialist nurseries grow it for the horticultural
peaty and water-logged and shallow over igneous trade. It is also grown in botanic gardens. Diselma
rock. Annual precipitation is very high, snow and archeri (the vernacular name is rarely used) is a
frost are possible year-round, with snow cover at the slow growing, more or less erect shrub, depending
highest altitudes lasting several months. on what shoots are being used for propagation by
cuttings, and very suitable for rock gardens. ‘Read
Dwarf ’ is an even slower growing cultivar with light
Conservation
green young foliage from a plant found on Mt. Read
This monotypic genus, though endemic to Tasma- in the north of Tasmania.
nia, is widely distributed in the highlands, where
Falcatifolium de Laub., J. Arnold Arbor. 50: 308. 1969. Type: Falcatifolium falciforme
(Parl.) de Laub. [Podocarpus falciformis Parl.] (Podocarpaceae).

Latin: falcis = sickle; folia = leaf. also exist and one should consult the descriptions
before concluding an identification.
Description
1a. Low shrubs. Adult leaves 0.6–1 cm long, 1.8–2
Dioecious evergreen shrubs or trees. Resin in bark mm wide (only known from one locality in
and leaves. Bark thin, with scattered lenticels, even- New Guinea) F. sleumeri
tually flaking. Branching in monopodial trees in 1b. Shrubs or (small) trees. Adult leaves normally 363
pseudo-whorls, more irregular in shrubs, foliage longer than 1 cm 2
branches terminating in loose buds formed by nar- 2a. Juvenile leaves narrowly linear-lanceolate, up
rowly triangular scale leaves. Leaves spirally insert- to 7 cm long, 1.2–1.5 mm wide. Trees to 20 m
ed, dimorphic, more or less appressed scale leaves tall F. angustum
on leading and fertile shoots, alternating with more 2b. Juvenile leaves usually shorter than 3 cm, if
or less distichously spreading, bilaterally flattened, longer (to 12 cm) then wider than 5 mm. Shrubs
obliquely lanceolate-falcate foliage leaves on (lat- or trees to 22 m tall 3
eral) vegetative shoots, latter leaves single-veined, 3a. Juvenile leaves nearly linear, 1–2 cm long, 1.5
much larger than scale leaves, but variable in size, mm wide, adult leaves 3–6 mm wide with ob-
with stomata on both sides (leaves amphistomatic). tuse or sometimes acute apex. Receptacle when
Pollen cones on axillary, usually solitary but some- full-grown 20 × 8 mm F. taxoides
times clustered, short, scaly shoots (peduncles), cy- 3b. Juvenile leaves falcate or linear-lanceolate, 2.5–
lindrical and catkin-like; microsporophylls small, 12 cm long, 3.5–12 mm wide, adult leaves (1.5–)
triangular or acuminate, with two globose pollen 2–9 mm wide with acuminate or pungent apex.
sacs containing bisaccate pollen. Seed cones solitary Receptacle when full-grown smaller than 12 × 6
on axillary, scaly dwarf shoots (peduncles), consist- mm 4
ing of several spirally arranged bracts; bracts becom- 4a. Juvenile leaves 10–12 cm long, 10–12 mm wide;
ing swollen, fleshy and red, forming an irregularly adult leaves falcate or S-curved F. falciforme
shaped receptacle, with one fertile bract at or near 4b. Juvenile leaves not longer than 7.5 cm and not
apex bearing an inverted ovule on the adaxial side. wider than 7 mm, adult leaves falcate (some-
Seeds at an oblique angle to axis of receptacle, erect, times at base only), not S-curved 5
surrounded at base by a swollen epimatium, ovoid 5a. Leaves flushing whitish/yellowish green or
but more or less flattened, with two lateral ridges. glaucous; juvenile leaves only slightly longer
than adult leaves, to 3 cm long. Pollen cones
6 species. 0.5–1.3 cm long F. papuanum
5b. Leaves flushing pinkish or purplish; juvenile
leaves markedly longer than adult leaves, to
Distribution
7.5 cm long (adult leaves to 4 cm long). Pollen
Malesia: Peninsular Malaysia, Borneo, Sulawesi, cones 5–6 cm long F. gruezoi
Phulippines, Maluku [Moloccas], New Guinea. SW
Pacific: New Caledonia.
Falcatifolium angustum de Laub., J. Arnold Arbor.
50: 312. 1969. Type: Malaysia: Sarawak, Bintulu, Sg.
Key to the species of Falcatifolium
Meluang W. S., E. F. W. Brunig S8866 (holotype L).
The reproductive organs (pollen cones and seed
cones) of some species remain not or poorly known;
Etymology
for this reason their use in this key remains tenta-
tive and should be treated with some caution. Tran- The species epithet refers to the narrow leaves.
sitional forms between juvenile and adult leaf forms
 Vernacular names Uses
No vernacular names are recorded for this species. No uses have been recorded of this rare species.

Description
Falcatifolium falciforme (Parl.) de Laub., J. Arnold
Trees to 20 m tall; trunk to 30 cm d.b.h. Bark Arbor. 50: 309. 1969. Podocarpus falciformis Parl.,
smooth, becoming rough and scaly, purplish brown, in Candolle, Prodr. 16 (2): 685. Type: Malaysia:
weathering grey. Foliage branchlets terete, glabrous. Sarawak, Gunung Pueh, (Mt. Poe), O. Beccari 2437
Juvenile leaves narrowly linear-lanceolate, curved (lectotype FI). Fig. 121, 122, 123
364 forward, up to 7 cm long and 1.2–1.5 mm wide above
a sessile or short petiolate, decurrent base, gradu- Falcatifolium usan-apuense de Laub. & Silba, Phyto-
ally tapering towards a pungent apex, flattened. logia 68: 32, 512. 1990.
Adult leaves shorter, 1.8–3.5 cm long and 1.5–2.5 mm
wide, decurrent at base, with nearly parallel sides for
Etymology
most of their length, flattened but keeled on both
sides forming an acutely raised midrib, tapering to a The species epithet (Latin falcis = sickle) refers to the
pungent apex. Stomata in several lines on both sides falcate leaves.
from base to apex (leaves amphistomatic). Pollen
cones terminal or lateral, axillary to leaves, in a more
Vernacular names
or less immature state 8 mm long and 2 mm diam.
Seed cones not observed. ekor sabit (Malay); kayu china (Sabah)

Taxonomic notes Description

This species is distinct in its extremely narrow leaves Shrubs or small trees 1.5–12 m tall, occasionally in
but the cones remain unknown or, in the case of forest a tall tree to 35–40 m, with an erect, mono­
male cones, poorly known. This taxon has been col- podial trunk to 80 cm d.b.h. and a spreading, more
lected from two locations near the coast in Sarawak or less open crown. Bark smooth, in large trees
(NW Borneo) namely Bintulu and Kuching and is becoming fissured and scaly, dark brown weather-
undoubtedly rare. ing grey-brown; inner bark slightly fibrous, grey-
ish brown, exuding red resin or sap in slash. Leaves
of two types: scale leaves and foliage leaves. Scale
Distribution
leaves on leading shoots and on the base of lateral
Borneo: Sarawak (two locations on coast). shoots, more or less appressed, subulate to narrowly
TDWG codes: 42 BOR-SR lanceolate, 4–6 × 1–2 mm, sometimes transforming
into small foliage leaves. Foliage leaves on seedlings
and saplings much longer than on mature plants,
Ecology
flushing leaves pinkish to purplish red, turning lus-
Falcatifolium angustum occurs in kerangas (forest trous green. Juvenile type leaves linear-lanceolate
on podzolised white sands) at 90–240 m altitude to more often falcate, on seedlings soon becoming
near the coast. The two populations occur in open large to 10–12 cm long, rapidly widening from a pet-
forest with among other trees Gymnostoma sp. (Cas- iole-like base to 10–12 mm well below the middle,
uarinaceae), Parastemon sp. (Chrysobalanaceae) and gradually tapering and curved forward to an acu-
Shorea albida (Dipterocarpaceae). minate apex, midribs on either side a thin, barely
raised ridge running from base to apex. Adult shade
leaves falcate or more commonly slightly S-curved,
Conservation
3–7 cm × 5–9 mm, tapering at both ends, midrib
IUCN: VU (D2) not or only slightly raised; apex acuminate. Adult
sun-exposed leaves still shorter, variable in size on a
branch, ­(0.6–)2–4 cm × (2–)4–8 mm, shortest ones
often more abrubtly tapering at both ends, other- dwarfed; here it forms shrubs or small trees to 12 m
wise of similar shape as shade leaves or sometimes tall. However, its altitudinal range is from 300 m to
less curved. Stomata in numerous intermittent lines 2100 m a.s.l. In kerangas (forest on leached podzol-
on both sides of leaves from base to apex. Pollen ic sands) it is an understorey tree or occasionally a
cones solitary or clustered, on axillary or sometimes canopy tree, with e.g. Agathis borneensis, Sundacar-
terminal scaly dwarf shoots, 2–4 cm × 2.5–3.5 mm; pus amarus, Nageia wallichiana and Dacrycarpus
microsporophylls terminating in an acuminate apex imbricatus as (co)dominant or emergent conifer
above two pollen sacs. Seed cones solitary on axil- trees. Occasionally it is found as a co-dominant or
lary dwarf shoots with acuminate scale leaves; re- more or less emergent large tree in lowland to sub-
ceptacles 4–6 mm long, swelling to 10–12 mm and montane primary rainforest on more fertile soils;
becoming succulent and orange or bright red. Ma- these individuals are scattered and rare and may be 365
ture seed single, ovoid, obliquely attached, 6–7 mm associated with episodal disturbance and succession
long, slightly flattened with two lateral ridges, green events, which initially led to more abundant coni-
turning purplish black. fers, most of which were in a later successional stage
replaced by angiosperms.
Taxonomic notes
Conservation
De Laubenfels (1988) in his treatment of the spe-
cies Falcatifolium falciforme for Flora Malesiana IUCN: LC
mentioned a specimen with very small leaves (6 × 2
mm) “in the form of shade leaves” collected on the
Uses
Usan Apua Plateau of Sarawak in “high kerangas”
from a tree said to be 24 m tall. He suggested that The rather rare large trees of this species will be
this might be a new species, whereupon Silba pro- logged together with other podocarps when grow-
ceeded to erect a new species F. usan-apuensis based ing outside protected areas. Its wood is traded as
on G. H. Pickels SAR-3925 (holotype in SAR; there ‘sempilor’ together with that of Dacrydium, Dacry-
is no material in FHO, another herbarium where carpus and Phyllocladus. Falcatifolium wood is light
Pickels deposited specimens) collected on this pla- and easy to work; it is used in light construction,
teau. Measurements of leaves given by Silba (6–19 doors, windows, joinery, furniture, interior finish-
× 2.5–4.5 mm) presumably from this specimen are ing, veneers as well as boat masts and crates. The
small, but in the lower range of the sun-exposed wood is not durable and therefore unsuitable for
leaves of the species F. falciforme and no evidence work that will be exposed to outdoor conditions. It
other than De Laubenfels’ remark is presented that is in cultivation only as specimens in some tropical
the leaves of the Pickels specimen are shade leaves. botanic gardens.
A 24 m tall tree would most likely have its foliage
in or above the forest canopy and not in the shade
of it. The two different kinds of leaves have no truly Falcatifolium gruezoi de Laub., Fl. Malesiana, ser.
distinct shapes but merge almost imperceptibly into 1, 10 (3): 373. 1988. Type: Philippines: Mindoro,
each other. Parabatugan, Mt. Halcon, [“Paitan access,
Paitaraan”], W. M. Gruezo 4052 (holotype L).
Distribution
Etymology
Borneo (including Lingga Island), Peninsular Ma-
laysia. This species was named after W. M. Gruezo, the col-
TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR lector of the type specimen.
MLY-PM
Vernacular names
Ecology
No vernacular names are recorded for this species.
Falcatifolium falciforme is most common on moun-
tain ridges where the forest is more sparse or
 collection which probably represents shade leaves,
Description
would be the most striking character of this species
Small trees 4–12 m tall, with a monopodial trunk to distinguishing it from F. falciforme as well as other
25 cm d.b.h. Bark smooth, becoming rough, brown species.
weathering grey-brown. Leaves of two types: scale
leaves and foliage leaves. Scale leaves on base of
Distribution
lateral shoots, more or less appressed, subulate to
narrowly lanceolate, 3–4 × 1 mm, sometimes trans- Malesia: Maluku [Moluccas] (Obi), Philippines, Su-
forming into small foliage leaves. Foliage leaves on lawesi.
seedlings and saplings much longer than on ma- TDWG codes: 42 MOL PHI SUL
366 ture plants, flushing leaves pinkish to purplish red,
turning green. Juvenile type leaves linear-lanceolate
Ecology
to falcate, on seedlings soon becoming large to 7.5
cm long, widening from a petiole-like base to 6–7 Falcatifolium gruezoi occurs in lower montane to
mm well below the middle, gradually tapering and montane rainforest. It is commonly associated with
curved forward to an acuminate apex; midribs on other conifers, e.g. Agathis dammara, Dacrydium
either side a thin, barely raised ridge running from spp., Nageia wallichiana, and Sundacarpus amarus.
base to apex. Adult shade leaves falcate, 20–40 × Angiosperms are often abundant but not dominant;
5–8 mm, tapering at both ends, midrib not or only dominance is usually attained by Agathis, Nageia, or
slightly raised; apex acuminate. Adult sun-exposed Sundacarpus on these sites, while F. gruezoi grows
leaves much smaller, 8–15(–20) × 2–4(–6) mm, of under canopy of these. On mountain ridges it can
similar shape as shade leaves, glaucous. Stomata in compete with angiosperms where these remain
numerous intermittent lines on both sides of leaves smaller trees.
from base to apex. Pollen cones solitary, on axillary
or sometimes terminal scaly dwarf shoots, 5–6 cm ×
Conservation
1.5–3 mm; microsporophylls terminating in an acu-
minate apex above two pollen sacs. Seed cones soli- IUCN: NT
tary on axillary dwarf shoots with acuminate scale
leaves, receptacle 2 mm long (larger when mature?);
Uses
mature seed single, ovoid, obliquely attached, 6–7
mm long, slightly flattened with two lateral ridges, This species remains a small tree and is therefore
dark brown. unlikely to be of value for its timber. No other uses
are known of it.
Taxonomic notes
This species was described by De Laubenfels Falcatifolium papuanum de Laub., J. Arnold Arbor.
based on a sterile collection with small, glaucous 50: 312. 1969. Dacrydium papuanum (de Laub.)
leaves from Mt. Halcon in the Philippines and on Whitmore, in Whitmore et al. (eds.), Tree Fl.
a description published in the Philippine Journal of Indonesia, Checkl. Irian Jaya: 233. 1997. Type: Papua
Science, Botany Vol. 6: 153–154 (1911) by F. W. Fox- New Guinea: Morobe District, Wau Subdistrict,
worthy under Dacrydium falciforme. The latter ac- Edie Creek, D. J. de Laubenfels P 483 (holotype A).
count refers to a collection by Elmer D. Merrill from
Mt. Halcon which had pollen cones ca. 6 cm long
Etymology
and 3 mm diam.; however, a duplicate at K of the
cited collection (Merrill 5744) is sterile. It is possi- The species epithet refers to its main distribution in
ble that a duplicate specimen in the Philippine Na- Papua New Guinea.
tional Herbarium (PNH) of this collection has such
pollen cones, but I have not seen it. If correct, its
Vernacular names
long, thread-like pollen cones, rather than the small,
glaucous leaves, which are not present on Merrills mungag, tugi (Papuan languages)
 Description
Conservation
Trees to 22 m tall, with an erect, monopodial trunk
to 40 cm d.b.h. and a more or less conical crown in Despite logging in areas within the distribution of
forest. Bark smooth, becoming fissured and scaly on this species which must have included it (no distinc-
the lower trunk, dark brown weathering grey; in- tion is being made and timber logs once deprived
ner bark slightly fibrous, red-brown. Leaves of two from foliage can only be assigned to ‘podocarp’ with
types: scale leaves and foliage leaves. Scale leaves on confidence), it is thought to grow in enough remote
leading shoots and on base of lateral shoots, more or and untouched forest localities to be as yet out of
less appressed, subulate to narrowly lanceolate, 2–5 risk of extinction.
× 1–2 mm, sometimes transforming into small foli- IUCN: LC 367
age leaves. First leaves on seedlings linear, spreading
at wide angles, petiolate, 10–15 × 1 mm, later leaves
Uses
on seedlings and small saplings slightly larger than
adult leaves, flushing leaves pale whitish green or The wood of this species provides good timber and
yellowish green to glaucous, becoming green with is being logged together with species of Dacry­carpus,
a glaucous underside. Adult shade leaves falcate at Dacrydium and Phyllocladus, between which the
base, then more or less straight, 10–22 × 2–4 mm, timber trade makes no distinction. Its relative rarity
more or less petiolate at base, tapering slightly in and usually small size make it unlikely that it will
the distal part, more or less coriaceous, midrib not contribute substantially to that trade. Aside from a
or only slightly raised; apex pungently acuminate. few specimens in greenhouses of botanical gardens
Adult sun-exposed leaves smaller, 8–13 × 1.5–2 mm, this species is not in cultivation.
narrower than shade leaves and sometimes more
curved. Stomata in numerous intermittent lines on
both sides of leaves from base to apex. Pollen cones Falcatifolium sleumeri de Laub. & Silba, Phytologia
solitary on axillary or sometimes terminal scaly 65: 329. 1988. Type: Indonesia: Papua, Arfak Mts.,
dwarf shoots, 0.5–1.3 cm × 2–2.5 mm; microsporo- Mt. Nettoti, P. van Royen & H. O. Sleumer 8203a
phylls terminating in an acuminate apex above two (holotype L).
reddish pollen sacs. Seed cones solitary on axillary
dwarf shoots with acuminate scale leaves; recepta-
Etymology
cles 7–10 mm long, swelling and becoming succulent
and bright red; mature seed single, ovoid, obliquely This species was named after Hermann Otto Sleum-
attached, 5–6 mm long, slightly flattened with two er (1906–1993), a German botanist who worked at
lateral ridges, light or dark brown. the Rijksherbarium in Leiden.

Distribution Vernacular names

New Guinea. No vernacular name is known for this rare species.
TDWG codes: 43 NWG-IJ NWG-PN
Description
Ecology
Low, more or less flattened or prostrate [?] shrubs
This species is locally common in montane rain­forest 20 cm tall or more [based on a single specimen de-
dominated by Fagaceae (Nothofagus, Litho­carpus), scribed to be of that dimension this habit and size
Myrtaceae, Cunoniaceae, and Podocarpaceae. On remain speculative]. Bark scaly, greyish brown.
mountain ridges it can reach the canopy, but in taller Branches numerous, spreading to erect, creating a
forest on moist slopes it is usually a sub-canopy tree, spreading, flattened crown covering the ground for
not exceeding 15 m in height. On exposed mountain several square meters. Leaves linear-ovate, curved at
summits this species is sometimes reduced to dwarf the shortly petiolate base, falcate or nearly straight,
size. 0.6–1 cm long and 1.8–2 mm wide, midrib raised on
 the abaxial side; margins slightly revolute, more or IUCN: NT
less abrubtly tapering to an apiculate apex with a
prickly spine 0.2 mm long. Reproductive structures
Uses
not known.
No uses are recorded of this species; if its habit is
indeed a prostrate shrub it could be of interest to
Taxonomic notes
horticulture, but only in countries with a frost-free
From Mt. Nettoti at 1920 m a.s.l. in the Vogelkop climate.
Peninsula a dwarfed plant was collected with very
small leaves 6–10 × 2 mm which are morphologi-
368 cally similar to leaves on trees of Falcatifolium pa- Falcatifolium taxoides (Brongn. & Gris) de Laub.,
puanum, in particular sun-exposed leaves. The J. Arnold Arbor. 50: 310. 1969. Dacrydium taxoides
leaves were illustrated with a small photograph in Brongn. & Gris, Ann. Sci. Nat. Bot., sér. 5, 6: 245.
the protologue of F. papuanum (De Laubenfels, 1866. Type: New Caledonia: Grande Terre, Province
1969: 313, fig. 6b) and it was speculated that it might Nord, Balade, [“montagnes de Balade”], E. Vieillard
represent a distinct “entity” or even a distinct species 1259 [1855–60] (lectotype P). Fig. 124
(De Laubenfels, 1988: 374). De Laubenfels & Silba
(op. cit.) proceeded to describe it as a new species,
Etymology
F. sleumeri, based on the same specimen already
mentioned in the account of 1969 and with no fur- The species epithet refers to Taxus, presumably from
ther knowledge or information, e.g. on reproduc- a resemblance of its leaves with those of yew.
tive structures; that single specimen, P. van Royen &
H. Sleumer 8203a (holotype L) is apparently sterile.
Vernacular names
This specimen is perhaps distinct in its decumbent
habit (shrub to 20 cm tall) and small leaves, but its No common names are known for this species.
reproductive structures remain unknown; therefore
its taxonomic status is not certain.
Description
Shrubs or small trees 2–15 m tall; trunk with max.
Distribution
d.b.h. 25 cm. Bark smooth becoming rough on larg-
New Guinea: Vogelkop Peninsula (Mt. Nettoti). er stems, brown weathering grey. Crown usually
TDWG codes: 43 NWG-IJ open and sparse in sub-canopy trees, more dense
in trees growing in open vegetation. Leaves of two
types: scale leaves and foliage leaves. Scale leaves on
Ecology
leading shoots and on base of lateral shoots, more
Stated to occur in “dark mossy forest”, so presum- or less appressed, subulate to narrowly lanceolate,
ably a ground covering species under a canopy of 2–3 × 1 mm, sometimes transforming into small fo-
trees. The altitude is given as 1920 m above sea level. liage leaves. First leaves on seedlings nearly linear,
spreading at wide angles, petiolate, 12–20 × 1.5 mm,
strongly keeled abaxially, soon replaced by adult
Conservation
leaves, flushing leaves pink with whitish bloom,
Known from only one locality and its type speci- becoming bright green on both sides. Adult leaves
men, but due to poor collecting records from the falcate at base, then more or less straight or curved
area nothing can be concluded from this as to rar- or ovate-oblong, variable in size on a single branch-
ity, population size etc. It is described as a prostrate let, 0.8–2.5 cm × 3–6 mm, more or less petiolate at
shrub in “dark, mossy forest”. The fact that is has base, tapering in the distal part; midrib raised on
been described in 1988 and no subsequent collec- both sides or inconspicuous; apex obtuse or some-
tions were made is undoubtedly due to its very local times acute. Stomata in numerous intermittent lines
occurrence. Rather than maintain it as DD the Con- on both sides of leaves from base to apex. Pollen
ifer Specialist Group believes this species should be cones on axillary or sometimes terminal scaly dwarf
flagged as NT as it is surely very rare and restricted. shoots, often 2–3 together, 1.5–2.5 cm × 1.5–2 mm;
microsporophylls terminating in an acuminate apex montana (on Mt. Panié) as well as angiosperm trees
above two yellow pollen sacs. Seed cones solitary on e.g. in the Meliaceae. Its most peculiar associate is
axillary dwarf shoots with acuminate scale leaves, the parasitic podocarpaceous conifer Parasitaxus
with 10–12 scales subtended by elongated bracts; usta, to which it is the only host. This diminutive
cone swelling to an irregularly shaped receptacle ca. shrub or treelet usually attaches to its roots and
20 × 8 mm and becoming succulent and bright red; seems to grow from the litter under Falcatifolium
mature seed single, ovoid, obliquely attached near taxoides. The large and succulent red receptacles are
apex, 6–7 × 3–4 mm, slightly flattened with two lat- conspicuous and eaten by birds, who disperse the
eral ridges, reddish maturing to dark brown. single seed thereby.

369
Distribution Conservation
New Caledonia: throughout Grande Terre. IUCN: LC
TDWG codes: 60 NWC
Uses
Ecology
This small tree is not exploited for its timber and
Falcatifolium taxoides is common as an understorey other uses in New Caledonia are unknown. It is
tree in montane humid forests from lowland hills in in cultivation only in some collections of botanic
the south of Grande Terre to the summit ridges of gardens; it could grow outside only in subtropical
the highest mountains on the island at 1400 m or to tropical climates. It has attractive pink flushing
even more. It occurs on ultramafic soil derived from leaves and larger pollen cones and red, succulent
serpentine as well as on acidic metamorphic rock receptacles than other species and would make an
(micaschist). It is associated with Araucaria mon- attractive small amenity tree in tropical countries.
tana, A. laubenfelsii, A. humboldtensis, and Agathis
 Fitzroya Lindl., J. Hort. Soc. London 6: 264. 1851. Type: Fitzroya cupressoides
(Molina) I. M. Johnst. [Pinus cupressoides Molina] (Cupressaceae).

Named after Robert FitzRoy (1805–1865), who current base; leaf colour lustrous green to glaucous
achieved lasting fame as the captain of HMS Beagle green, with whitish wax on stomatal zones. Pollen
during Charles Darwin’s famous voyage. cones predominantly terminal, solitary, cylindrical,
6–8 × 2–3 mm; microsporophylls 15–24, in alternate
whorls of 3, imbricate, peltate-ovate with acute apex,
Description
keeled abaxially; margins entire, bearing (2–)4–6
370 See the species description. small pollen sacs on abaxial side. Seed cones termi-
nal or subterminal, formed by 2–3 whorls of slightly
modified scale leaves, followed by 2 alternate whorls
Distribution
of 3 fertile scales, maturing in 1 season to cones with
As for the species. wide spreading scales 10–14 mm diam. Bract-scale
complexes thin woody, obovoid-spathulate, largest
one ca. 5 × 4 mm, with thickened, wrinkled api-
Fitzroya cupressoides (Molina) I. M. Johnst., Contr. cal part and subapical, curved umbo of exserting
Gray Herb., n. s. 70: 91. 1924. Pinus cupressoides bract apex 1–2 mm long. Columella variably shaped,
Molina, Sag. Stor. Nat. Chili: 168. 1782. Type: Chile: trigonal to tripartite, ca. 4 mm long, flattened, not
Los Lagos, Cordillera de la Costa, 30 km from San woody, resinous-glandular, more or less translucent.
Juan de la Costa, J. L. Morrison 17636 (neotype K). Seeds usually only well developed on upper fertile
Fig. 125, 126 whorl, 3–4 × 2 mm, ovoid-oblong, flattened, with a
concave hilum; wings usually 2, sometimes 3, cres-
cent-shaped with undulate margins, ca. 2 mm wide.
Etymology
The species epithet means “like a cypress” (Cupres-
Distribution
sus).
S Argentina: Neuquén, Rio Negro; S Chile: Los La-
gos. For the most part, the still existing populations
Vernacular names
of Fitzroya cupressoides are in Chile, now disjunct in
Patagonian cypress, Alerce (Spanish), Lahuán the coastal Cordillera, some in the Central Valley,
and others in the Andes, where they cross the bor-
der into Argentina.
Description
TDWG codes: 85 AGS-NE AGS-RN CLS-LL
Trees to 50(–60) m tall, evergreen, dioecious, rarely
monoecious; trunk monopodial, straight, buttressed
Ecology
in old individuals and often with root suckers, to 3–5
m diam. above buttress. Bark on trunk thick, with A dominant or emergent tree usually forming more
deep fissures, exfoliating in long strips, reddish or less large stands (‘alerzales’) scattered in the Val-
brown. Branches spreading horizontally or curved divian evergreen rainforest. Understorey tree species
down, forming a conical or pyramidal crown. Foliage are e.g. Podocarpus nubigenus, P. salignus, Prumno-
branches spreading or pendulous, ultimate branch- pitys andina, Saxegothaea conspicua, Pilgero­dendron
lets 1–4 cm long, more or less terete, densely covered uviferum, Nothofagus betuloides, N. dombeyi, N. nit-
with leaves, persistent. Leaves scale-like, in alternate ida, N. obliqua, Drimys winteri, and Laurelia philip-
near-whorls of 3, imbricate, decurrent, with distal piana. There are various forest types with shifting
part variously (re)curved; apex incurved but free, dominance of these associates on altitudinal gradi-
lanceolate to ovate, 4–6 mm long on younger trees, ents and on soil types, with e.g. a very wet, acidic
2–3 mm long on older trees, 1–2 mm wide, keeled woodland bog type at middle to high (–1200 m)
abaxially, adaxially with a midrib; margins entire; elevation dominated by Pilgerodendron and Fitz­
apex obtuse; stomata abaxially in a few lines on de- roya and a shrublayer of the conifer ­Lepidothamnus
fonkii in carpets of Sphagnum. Another forest type populations of Fitzroya are protected is The Pumalin
occurs between 800–1200 m on better drained Park in southern Chile which was set up in 1991 to
volcanic soils with dominance of Nothofagus betu- help protect 300,000 ha of pristine rainforest, mak-
loides and emergent Fitzroya; it is there dependent ing the largest private park in the world. In the late
on episodal disturbance in the absence of which 1990s, studies carried out into the genetic variability
Notho­fagus would prevent its regeneration. Fitzroya of Fitzroya using DNA markers have helped guide
cupressoides is one of the longest living trees in the researchers from the Forestry Faculty of the Univer-
world, with record ages counted between 3500 and sidad Austral de Chile in establishing an important
4000 years. The soils on which Fitzroya occurs are restoration programme in the Central Depression.
usually poorly drained and water-logged, or incipi- IUCN: EN (A1c-d, A2c-d)
ent soils on volcanic ash deposits, at lower elevations 371
sandy or loamy soils predominate, often with a very
Uses
high organic content and a low pH (4.0–5.1) in the
upper layer. Mean annual precipitation ranges from This species has been extensively used for building
2000–6000 mm, mostly as winter rains, with snow timber and especially shingles for roofing. Its ex-
at the higher elevations. tremely rot resistant wood is so durable, that shin-
gles on roofs 130–150 years old are still perfectly
sound (Golte, 1996). In 1973 logging was still going
Conservation
on and sawmills were well stocked and busy, but
The range of Fitzroya cupressoides has historically exploitation has declined sharply since about 1980,
been reduced greatly by centuries of logging, prob- with export no longer legally possible (an illegal
ably aggravated by recurring fires both natural and shipment of wood was intercepted in the UK and
man-made. It has largely disappeared from the low- given to the Royal Botanic Gardens, Kew, who built
land areas N of Puerto Montt, where it was once ex- an education centre from it at its garden Wakehurst
tensive and abundant (Golte, 1996). It is not doing Place in Sussex, England). This conifer was intro-
well in many of the remaining, mostly upland and duced to cultivation by William Lobb in 1847 (possi-
very wet stands, and large tracts of land merely have bly based on his brother’s collection) and it remains
standing dead snags with little or no natural regen- an uncommon tree mostly confined to dendrologi-
eration. Exploitation has been on a devastating scale cal collections. Recently renewed seed collecting has
with no intent of sustainability until 1976, when replenished the original introduction, which ap-
logging of living trees was banned by the Chilean peared to have been propagated from cuttings; new
government and the species was declared a National plants have been distributed in the UK largely as ex
Monument. Fires continue to kill trees, which can situ conservation stock by the International Coni-
then be logged and the land converted to pasture. fer Conservation Programme of the Royal Botanic
Since 1973 the species is listed on CITES Appendix Garden at Edinburgh, Scotland. As a CITES Ap-
I which prohibits all export. Although it is afforded pendix I listed species its commercial distribution
protection in both Argentina and Chile in National will remain restricted and plants can only be sold if
Parks, most populations occur on private property. they originate from legal stock in cultivation outside
The most notable private property where important Chile and Argentina.
 Fokienia A. Henry & H. H. Thomas, Gard. Chron., ser. 3, 49: 67. 1911. Type:
Fokienia hodginsii (Dunn) A. Henry & H. H. Thomas [Cupressus hodginsii Dunn]
(Cupressaceae).

Fokien is an earlier European spelling for Fujian, a leaves appressed, obtuse, variable in size from 2–7
province of China. × 1–2 mm on ultimate branchlets; stomata in two
bands separated by a midrib. Lateral leaves longer
than facials or nearly equal in length, bilaterally
Description
flattened, variable, 4–10 × 2–3(–4) mm on ultimate
372 See the species description. branchlets; apex incurved, free to appressed, acute
to obtuse; stomata in a single whitish band, eglandu-
lar. Pollen cones terminal, 4–5 × 1.5–2 mm, initially
Distribution
subglobose but elongating when maturing, with 10–
As for the species. 12 decussate, broadly ovate-acute microsporophylls
each bearing 3 small, abaxial pollen sacs. Seed cones
terminal, maturing in the second year, caducous, to
Fokienia hodginsii (Dunn) A. Henry & H. H. Tho- 15–25 × 12–22 mm, subglobose, brown. Bract-scale
mas, Gard. Chron., ser. 3, 49: 66–68, f. 32–35. 1911. complexes (10–)12–16, decussate, spreading at ma-
Cupressus hodginsii Dunn, J. Linn. Soc., Bot. 38: 367. turity, to 10 × 16 mm (uppermost 2–4 much smaller
1908; Chamaecyparis hodginsii (Dunn) Rushforth, and sterile), cuneate-peltate, flattened, woody, with
J. Biol. (Vietnam) 29 (3): 38. 2007. Type: China: Fu- irregular margins, distally with a central depression
jian, Min Jiang, Nanping, [“Yenping”], A. E. N. Hod- and a small mucronate umbo (bract tip), dull brown,
gins HK 3505 (holotype K). Pl. 14, Fig. 127 with striated anterior part, orange- to reddish brown
with whitish seed marks. Seeds 20–30 per cone, 4–5
mm long, ovoid and 3–4 ridged, with acute apex and
Etymology
truncate base (hilum), brown, wings 2, very unequal
The species epithet is after A. E. N. Hodgins, who in size and shape; larger wing ca. 6 × 5 mm, smaller
collected it in 1905. one ca. 1.5 mm, or a mere strip near the seed apex.

Vernacular names
Taxonomic notes
Fujian Cypress; fu jian bai (Chinese)
Some authors have placed this species either in Cu-
pressus or (recently) in Chamaecyparis based on
Description
comparisons of (some) gross-morphological char-
Trees to 30 m tall, evergreen, monoecious; trunk acters. Detailed analyses of the ontogeny of the seed
monopodial, to 1 m d.b.h. Bark on trunk nearly cones (Jagel, 2001; Farjon, 2005a) and phylogenetic
smooth, becoming irregularly fissured and flaking, analyses based on morphology (Farjon, 2005a) as
purplish brown. Branches long, spreading or down- well as DNA sequence data (Gadek & Quinn, 1993)
curved, self-pruning lower down the trunk, forming have demonstrated that Fokienia hodginsii represents
a broadly pyramidal, in old trees rounded and dense an evolutionary lineage separate from these two gen-
crown. Foliage branches spreading in flattened era and therefore merits recognition at the rank of
sprays (plagiotropic), ultimate branchlets pinnately genus. Similarities observed, especially in the foliage
arranged, forming frond-like sprays, but in older and in later (mature) phases of cone development,
trees of irregular and shorter length forming flat, are likely to be the result of convergent evolution.
rounded sprays, deciduous. Leaves decussate, im-
bricate, decurrent, scale-like, strongly dimorphic on
Distribution
flattened ultimate branchlets but increasingly less so
on older (whip) shoots, becoming long decurrent to China: Chongqing, Fujian, N Guangdong, Guangxi,
15 mm, with facials usually smaller and/or narrower Guizhou, S Hunan, W Jiangxi, SE Sichuan, SE Yun-
than the laterals, green or glaucous green. Facial nan, S Zhejiang; N Laos; Vietnam.
 6

373

1
2

Pl ate 1 4. Fokienia hodginsii. 1. Habit of tree. 2. Branch with foliage. 3. Branchlet with juvenile leaves
(lower side). 4. Branchlet with adult leaves. 5. Pollen cones. 6. Microsporophyll with pollen sacs. 7. Seed
cones. 8, 9. Seeds.
 TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- situation in China is possibly similar in many areas
SC CHC-YN CHS-FJ CHS-GD CHS-GX CHS-JX CHS- where mature forests are shrinking under popula-
ZJ 41 LAO VIE tion pressures. However, the species is very widely
distributed and regeneration has been observed in
most populations. Continued logging at current or
Ecology
increased rates may cause scarcity of large trees in
This conifer is usually a minor constituent of sub- the near future, but provided that sufficient forests
tropical to warm temperate evergreen (mixed) remain across its range, the species seems not to be
mesophytic forest, which in undisturbed state is at significant risk of becoming extinct.
dominated by numerous angiosperm trees e.g. IUCN: NT
374 Castanopsis spp., Quercus spp., Lithocarpus spp.,
Nyssa sinensis, Pasania spp., Schima argentea, S. su-
Uses
perba, and a few other conifers e.g. Cephalotaxus
fortunei and Nothotsuga longibracteata. With contin- The wood of this species, like other cupressaceous
ued disturbance conifers like Cunninghamia lanceo- trees in eastern Asia, is much valued for construc-
lata and especially Pinus massoniana or P. densata tion. In most areas where it occurs, wild growing
become more abundant and eventually dominate. trees are logged; its slow growth makes it a tree dif-
The altitudinal range of Fokienia hodginsii is from ficult to exploit economically in plantation forestry.
350 m to 2000 m a.s.l. Soils are acidic (pH 5–6) yel- In southern China and in Vietnam the rot resistant
low or brown loamy sands over sandstone, shales, and even-grained wood is prized for doors, window
granite, or rhyolite and well drained. Mean annual frames, and carved panels, as well as furniture. In re-
precipitation is 1200 mm or more. mote regions pit-sawn timbers are often transported
off the mountains by man-power to fetch high prizes
with carpenters in the towns. This conifer is cultivat-
Conservation
ed in cemeteries and parks in China, but rare else-
Mature trees are selectively logged in Vietnam and where despite its relative hardiness and suitability in
Laos, even from forest reserves in remote areas us- parts of the world with similar climates such as New
ing helicopters and possibly beyond sustainabil- Zealand, Atlantic SW Europe, northern California
ity, because the dark red timber is considered very and Oregon and southern Japan.
valuable (roughly estimated at US$ 2500/m3). The
Glyptostrobus Endl., Syn. Conif.: 69. 1847. Type: Glyptostrobus pensilis (Staunton ex
D. Don) K. Koch [Thuja pensilis Staunton ex D. Don] (Cupressaceae).

Greek: glypto- = cut into; strobus = cone; referring to leaves on ‘long’ shoots decurrent, imbricate, 1.5–3 ×
the margins of the seed cone scales. 0.5–0.8 mm, lanceolate, convex or keeled abaxially;
apex incurved but free, obtuse. Needle-like leaves
on ‘short’ shoots of two types, helically arranged,
Description
decurrent, and either more or less distichous, linear,
See the species description. 15–30 × 1.5–3 mm, bilaterally flattened, with entire
margins, acute, or more commonly in ranks of 3 or 375
nearly radially spreading, subulate, 4–10 × 0.4–0.6
Distribution
mm, abaxially keeled and adaxially grooved. Sto-
As for the species. mata on all leaves types on both sides, but more
on adaxial side. Pollen cones terminal, solitary, 3–5
mm long, subglobose to ellipsoid, with 15–20 heli-
Glyptostrobus pensilis (Staunton ex D. Don) cally arranged, imbricate, ovate-convex microspo-
K. Koch, Dendrol. 2 (2): 191. 1873. Thuja pensilis rophylls bearing (2–)4–6(–9) abaxial, small pollen
Staunton ex D. Don, in Lambert, Descr. Pinus, ed. sacs. Seed cones terminal, solitary on determinate
2, 2: 115. 1828. Type: China: Guangdong, [“Province branchlets, subtended by a few short, broad, keeled
of Guangton”], G. L. Staunton s.n. (holotype BM). leaves grading into bract-scale complexes, maturing
Fig. 128 in two seasons; mature cones pyriform to obovate,
14–25 × 10–15 mm, with 20–25 helically arranged,
imbricate woody scales, maturing brown. Bract-
Etymology
scale complexes increasing in size to 13 × 6.5 mm,
The etymology of the species epithet is unclear, obovate-­oblong, cuneate proximally, rounded dis-
perhaps it derives from Latin penicillatus = pencil- tally, connate but parting slightly at maturity, with
shaped [the branchlets]. 4–8(–11) subapical tooth-like lobes ca. 1 mm long
(often worn off) along distal margin and a (sub)cen-
tral, acute bract apex 2–3 × 2–3 mm; abaxial surface
Vernacular names
grooved or rough; adaxial surface with two unequal,
Chinese Swamp Cypress, Chinese Water Fir; shui deep seed cavities. Ovules 2 in axil of a fertile bract,
song (Chinese) erect, in well developed cones maturing to 20–30
seeds; seeds elliptic, 5–7 × 3–4 mm, slightly flat-
tened, often grooved, brown, with a single proximal,
Description
oblong and thin wing of 4–7 × 2–4 mm.
Trees to 15(–25) m tall (one planted tree in China
is 39 m tall), semi-evergreen, monoecious; trunk
Taxonomic notes
markedly buttressed at base especially when grow-
ing in watery substrate, to 1.2–1.5 m d.b.h. Bark on In many descriptive works on conifers, the name
trunk thin with long, irregular strips, whitish brown Glyptostrobus lineatus (Poir.) Druce has been ap-
or grey-brown. Branches long, slender, spread- plied to this taxon. That combination by Druce,
ing horizontally or curved down in old trees, foli- published in 1916, was based on Thuja lineata Poir.,
age in distinct tufts at ends of branches, forming a a species newly described in Supplement Vol. 5 of
pyramidal or more rounded, open crown. Foliage Lamarck’s famous Encyclopédie methodique (Bota-
branches alternate, spreading at less than 45 degrees, nique) in 1817. This description was based on a cul-
variable in length, dimorphic, ‘long’ shoots persist- tivated plant in the garden of “Monsieur Noisette”
ent, slender and flexible, ‘short’ shoots from termi- undoubtedly in France. Needless to say, this plant
nal or dormant lateral buds, deciduous, all covered cannot be traced and there is no specimen in the
with leaves. Leaves alternate to helically arranged, of Paris Herbarium (P) that can be linked to it. The de-
three types, all green with white stomatal dots. Scale scription fits Taxodium distichum var. imbricarium
 and not Glyptostrobus pensilis and while the former species was widespread in the Tertiary and may well
had been introduced to Europe and taken into culti- have occurred more widely along upland streams.
vation for more than two hundred years in 1817, the
latter has been introduced first in England “about a
Conservation
hundred years ago, but rarely endured our climate…
and seems to be unknown in the living state on the The present natural distribution of this species in
continent” (Henry & McIntyre, 1926). The plant in China is possibly restricted to several isolated locali-
Mr Noisette’s garden could not have been a speci- ties along upland streams in three or four provinces
men of Glyptostrobus. of SE China. However, it has been widely planted
along water courses in lowland S China for a very
376 long time and it may now be impossible to distin-
Distribution
guish between regional introductions and original
SE China: Fujian, S Hunan, Guangdong, Guangxi, occurrence, as many trees are found in ancient culti-
Jiangxi; introduced on Hainan Island and in Sichuan, vated landscapes. In N Vietnam it formerly occurred
possibly natural in SE Yunnan; extinct in the wild in along the Hong (Red) River but no natural stands
N Vietnam. This species has also been reported from are to be found there at present. Habitat change due
S Vietnam. The trees in Buon Uik and Krong Buc to intensive agriculture is the main cause of its de-
(Dac Lac Province) have been found at ca. 500–700 cline. In S Vietnam two small populations are prob-
m a.s.l. and were growing in “swampy forest or in ably indigenous and its status there is CR.
basaltic terrain.” This disjunct location and type of IUCN: EN (A1c; A2c; B1; B2a;C)
habitat do not fit well with what is known about the
occurrence in S China in river deltas just above sea
Uses
level; however, its indigenous status in S Vietnam
has recently been verified ecologically (Nguyen Tien The rather soft, yellowish wood is like most cupres-
Hiep et al., 2004). Recently, it has also been reported saceous wood decay resistant and finds uses in Chi-
from Khammouan Province in Laos. na ranging from furniture to building of bridges. The
TDWG codes: 36 CHS-FJ CHS-GD 41 LAO VIE wood of the roots is very light and due to its buoyan-
cy it is used in China to make life-saving rings. This
species is widely cultivated in southern China and
Ecology
planted along rivers and canals as well as in parks;
On river floodplains and in deltas, always near or in except for the latter localities mostly to harvest the
water, where it develops a buttressed base and oc- timber followed by replanting. This conifer is a truly
casionally pneumatophores; also extensively plant- semi-aquatic tree requiring permanent water logged
ed along rivers and canals. A heliophilous species, soil to thrive; planting it in rather than adjacent to a
intolerant of competition and usually growing in pond enhances its chances of survival. It can easily
pure stands or solitary along streams. It is restricted survive mild frosts in this habitat. It can be grafted
to the subtropical to tropical coastal lowlands, but onto rootstock of Taxodium, another conifer that
avoids saline and alkaloid soils or water. A probably grows well standing in water (but equally outside it).
indigenous occurrence in Vietnam along a marshy Its autumnal foliage colour is a deeper reddish than
small river at greater altitude (around 700 m) could that of Taxodium and it retains its leaves longer.
mean that it also occurs along inland waters. This
Halocarpus Quinn, Austral. J. Bot. 30 (3): 317. 1982. Type: Halocarpus bidwillii
(Hook. f. ex Kirk) Quinn [Dacrydium bidwillii Hook. f. ex Kirk] (Podocarpaceae).

Greek: halos = the sea; karpos = fruit. 2b. Trees to 25 m tall. Juvenile leaves 20–45 mm
long, 2–4 mm wide. Branchlets with adult scale
leaves 1.5–3 mm thick, nearly terete H. kirkii
Description
Dioecious, evergreen shrubs or trees. Resin ducts (1)
in leaves; resin in bark and leaves. Bark thin, hard, Halocarpus bidwillii (Hook. f. ex Kirk) Quinn,
with or without lenticels, becoming scaly and exfo- Austral. J. Bot. 30 (3): 317. 1982. Dacrydium bidwillii 377
liating in thin flakes. Branching variable, in shrubs Hook. f. ex Kirk, Trans. New Zealand Inst. 10:
often layering, irregular and ascending in trees. 388. 1878. Type: New Zealand: South Island,
Terminal buds on shoots with juvenile leaves only. Nelson, J. C. Bidwill 130 (lectotype K, sheet 1, here
Leaves spirally inserted, dimorphic, radially spread- designated). Fig. 129, 130
ing lanceolate-linear or linear leaves alternating
with appressed, rhombic scale leaves, amphistomat-
Etymology
ic. Pollen cones solitary or with 2–3 together at apex
of scale-leaved branchlets, small, with a few spirally The species epithet commemorates John Carne Bid-
arranged, reddish microsporophylls and two glo- will (1815–1853) an early collector of plants in New
bose, yellow pollen sacs containing bisaccate pollen. Zealand.
Seed cones solitary at apex of scale-leaved branch-
lets, consisting of a few slightly enlarged, spreading
Vernacular names
bracts continuing from and similar to the scale leaves
and distinguished by a reddish colour from normal Bog pine, Mountain pine, Tarwood
leaves; one or more fertile with 1 inverted ovule on
the adaxial side. Seeds 1–5 per cone, becoming erect,
Description
at base surrounded by the swollen epimatium form-
ing a white or yellow collar; seed mostly protrud- Prostrate shrubs, densely branched erect shrubs,
ing, more or less ovoid but flattened, with two lateral or sometimes a dwarfed tree to 4 m tall and 35 cm
ridges, striated, maturing to lustrous black. stem diam., commonly forming extensive thickets
by layering. Bark becoming rough, hard and greying
3 species. with age. Leaves of two distinct types: juvenile and
adult. Juvenile leaves on seedlings and young plants,
spreading radially, linear, (3–)5–10 mm long, 0.8–1.4
Distribution
mm wide, convex adaxially, concave abaxially, with
New Zealand. a midrib on that side; apex obtuse. Adult leaves
abruptly appearing above juvenile leaves, scale-
like, arranged in high spirals, on ultimate branch-
Key to the species of Halocarpus
lets seemingly decussate, completely clasping the
1a. Prostrate to erect shrubs, sometimes a dwarfed branchlets (including leaves 1.5–2 mm thick), rhom-
tree to 4 m tall. Juvenile leaves on seedlings and bic in appearance, 1.5 × 1.2 mm (larger on thicker
young plants. Swollen epimatium at the base of branches), weakly keeled and obtuse. Stomata of ju-
the seed white H. bidwillii venile leaves on both sides (leaves amphistomatic)
1b. Erect shrubs or trees to 25 m tall. Juvenile leaves but in more conspicuous bands on either side of
on seedlings, shrubs and trees. Swollen epima- the midrib on the adaxial side, on adult leaves scat-
tium at the base of the seed yellow or orange 2 tered to just below apex. Pollen cones often numer-
2a. Erect, dense shrubs or small trees to 10 m tall. ous, terminal on branchlets with adult leaves, 3–5
Juvenile leaves (5–)8–18 mm long, 1–1.5 mm mm long, slightly wider than branchlet; microspo-
wide. Branchlets with adult scale leaves 2–3 rophylls similar to scale leaves, yellow with 2–4 red
mm thick, angular H. biformis pollen sacs. Seed cones subterminal and erect on
 branchlets with adult leaves, solitary, consisting of
Conservation
4–5 leaf-like, slightly swollen bracts; one or some-
times two bracts fertile; epimatium aril-like, swollen IUCN: LC
and fleshy with irregular but clefted upper margin,
white, subtending and partially enclosing seed; seed
Uses
3–4 mm long, striated and crested, ripening from
green to black. This attractive conifer shrub remains exceedingly
rare in cultivation. In New Zealand there is a live-
ly gardening tradition but New Zealand gardens
Taxonomic notes
are very much Europe (read: England) orientated,
378 Brian P. J. Molloy annotated a specimen (sheet 1, with the native flora largely neglected. In England
a) at K collected by J. C. Bidwill under his number a few plants exist in arboreta, where it proves per-
130 as lectotype and specimens with number 131 as fectly hardy. It is a conifer shrub “worthy of more
“lectoparatype” of Dacrydium bidwillii Kirk in Sep- widespread trial” as Keith Rushforth noted 20 years
tember 1991. Lectotypification is only effected when ago in his book Conifers (Rushforth, 1987). Nothing
published, and this had not happened; in order to much seems to have happened and a reason for this
establish Molloy’s choice (but extend it to all four is perhaps that the horticultural trade is not interest-
branchlets on herbarium sheet 1) J. C. Bidwill 130 ed in ‘new’ species, but in new shapes and colour ‘ef-
on sheet 1 at K (specimens a-d which are a single fects’ which are more easily achieved working with
gathering from a single tree) is here designated as the well known species already in the trade.
the lectotype of Dacrydium bidwillii Kirk. Bidwill
130 (e-f) and 131 on sheet 2 at K were annotated by
Molloy as “lectoparatypes” and represent the resi- Halocarpus biformis (Hook.) Quinn, Austral.
due of original material at K after this choice; they J. Bot. 30 (3): 318. 1982. Podocarpus biformis Hook.,
are syntypes which no longer have a type status for Icon. Pl., n.s., 2: t. 544. 1843; Dacrydium biforme
purposes of nomenclature after a lectotype has been (Hook.) Pilg., in Engler, Pflanzenr. IV.5 [18]: 45.
chosen and published. 1903. Type: New Zealand: South Island, Southland,
Dusky Sound, A. Menzies s.n. (lectotype K). Fig. 131
Distribution
Etymology
New Zealand: North Island, South Island, Stewart
Island. The species epithet refers to the two distinct leaf
TDWG codes: 51 NZS types often encountered even on mature trees.

Ecology Vernacular names

Halocarpus bidwillii can form extensive thickets by Yellow pine, Pink pine, Alpine tarwood
layering on open, stony ground or in boggy grass-
land. In the northern parts of its range it is a mon-
Description
tane to subalpine shrub occurring between 600 and
1500 m a.s.l. but in the far south on Stewart Island Densely branched erect shrubs, or sometimes
it comes down to near sea level. It can be associated a small tree to 10 m tall and 60 cm d.b.h., with a
with H. biformis, Podocarpus nivalis, Phyllocladus rounded or flat-topped crown. Bark becoming
trichomanoides var. alpinus, Gaultheria depressa, rough, hard, scaly and greying with age. Leaves of
and Cassinia vauvilliersii in the central part of North two distinct types: juvenile and adult. Juvenile leaves
Island in similar open habitats, or with podocarp on seedlings to mature plants, spreading radially,
trees like Prumnopitys taxifolia, P. ferruginea and linear, (5–)8–18 mm long, 1–1.5 mm wide, adaxi-
Manoao colensoi in open coniferous forest at lower ally flat, abaxially with slightly revolute margins
altitudes further south. and a midrib; apex obtuse or apiculate. Adult leaves
abruptly appearing above juvenile leaves (or branch-
Uses
lets reverting to juvenile leaves), scale-like, spirally
arranged, completely clasping branchlets (including The pink-coloured wood with a pleasant, sweet fra-
leaves 2–3 mm thick and angular), rhombic in ap- grance is dense, fine-grained and extremely dura-
pearance, 1.5 × 1.2 mm (larger on thicker branches), ble. It is therefore suitable for outdoor applications
strongly keeled and obtuse. Stomata of juvenile of carpentry like garden furniture, but its generally
leaves on both sides (leaves amphistomatic) but in small size and often shrubby habit ensured that,
more conspicuous bands on either side of midrib even in the past when natural forest was much more
on adaxial side, on adult leaves conspicuous, scat- widespread than now, it was not a very common
tered to just below apex. Pollen cones often numer- commodity. It was used for railway sleepers, fence
ous, solitary or 2–3 together, terminal on branchlets posts and some outdoor building applications. State 379
with adult leaves, 3–4 mm long, slightly wider than protection of what is left of native trees prohibits
branchlet; microsporophylls similar to scale leaves, felling and so there is no commercial use of its wood
reddish, turning black, with 2–3 yellow pollen sacs. at present. It is not in cultivation outside a few bo-
Seed cones subterminal and erect on branchlets tanic gardens and arboreta.
having adult leaves tinged with red below the cones,
solitary, consisting of 4–5 leaf-like, slightly swollen
bracts; one or sometimes several bracts fertile; epi- Halocarpus kirkii (F. Muell. ex Parl.) Quinn,
matium aril-like, swollen and fleshy with irregular Austral. J. Bot. 30 (3): 318. 1982. Dacrydium kirkii
but clefted upper margin, yellow, subtending and F. Muell. ex Parl., in Candolle, Prodr. 16 (2):
partially enclosing seed; seed 2.5–3 mm long, stri- 495. 1868. Type: New Zealand: [“Great Barrier
ated and crested, ripening from green to black. Land(?)”], T. Kirk 81 (holotype not located, iso­
type K).
Distribution
Etymology
New Zealand: North Island, South Island, Stewart
Island. This species was named after the New Zealand bota-
TDWG codes: 51 NZN NZS nist Thomas Kirk (1828–1898), who studied the for-
est flora of that country.
Ecology
Vernacular names
Halocarpus biformis occurs more often in forested
habitats than H. bidwillii, but in boggy terrain in manoao (Maori)
the central volcanic plateau of North Island the two
may occur together. The altitudinal range of this
Description
species is between 550 m to 1400 m a.s.l. As a small
tree in forest it usually remains below the canopy of Trees to 25 m tall; trunk to 1 m d.b.h., short with
other trees. In open terrain it forms dense, rounded large, ascending branches forming a rounded or
shrubs to 2 m tall, surrounded by sedges and grasses flat-topped crown. Bark smooth with horizontal
(e.g. Deschampsia gracillima) indicating acid soil lenticels on young trees, becoming hard, scaly and
and water logging. Here it can be associated with the weathering grey, with exfoliating flakes covered in
conifers H. bidwillii, Podocarpus nivalis, and Phyl- small pustules exposing reddish brown bark. Leaves
locladus trichomanoides var. alpinus, as well as with of two distinct types: juvenile and adult. Juvenile
Cassinia vauvilliersii, Dracophyllum, Olearia colen- leaves on seedlings to trees up to 10 m tall, spreading
soi, and in forest edges Nothofagus solandri var. clif- radially, linear, 20–45 mm long, 2–4 mm wide, peti-
fortioides. olate and twisted at base, flat adaxially, with slightly
revolute margins abaxially, with a faint midrib on ei-
ther side; apex obtuse. Adult leaves abruptly appear-
Conservation
ing above juvenile leaves or on separate branches,
IUCN: LC scale-like, arranged in high spirals and on ultimate
branchlets seemingly decussate, completely ­clasping
 branchlets (including leaves 1.5–3 mm thick and making up the main canopy below them. Tall tree
nearly terete), rhombic in appearance, 1.2–1.5 × 1.5 ferns (­Cyathea, Dicksonia) often are abundant and
mm (larger on thicker branches), strongly keeled can reach the canopy as well. These are rainforests
and obtuse. Stomata of juvenile leaves on both sides with abundant epiphytes in the larger trees and nu-
(leaves amphistomatic) but more densely in bands merous smaller trees and shrubs in the understorey.
on either side of midrib on adaxial side, on adult
leaves conspicuous, scattered to just below apex.
Conservation
Pollen cones often numerous, solitary or 2–3 togeth-
er, terminal on branchlets with adult leaves, 3–4 mm This is the only species in the genus to attain large
long, slightly wider than branchlet; microsporo­ tree size and consequently it has been subject to
380 phylls similar to scale leaves, green turning yellow, overexploitation, with many of the forests in which
with 4–6 reddish pollen sacs. Seed cones subtermi- it occurred now gone. It has a limited natural dis-
nal and erect on branchlets with adult leaves tinged tribution and populations are now fewer and more
with red below cones, solitary, consisting of 4–5 leaf- scattered than before logging began in the 19th cen-
like, slightly swollen bracts; one or sometimes two tury. The reduction that occurred is difficult to esti-
or three bracts fertile; epimatium aril-like, swollen mate, but is likely to have been in excess of 50% from
and fleshy with irregular but clefted upper margin, its original area of occupancy (AOO), while podo-
orange, subtending and partially enclosing a seed; carp wood was being logged preferentially where
seed 3–4 mm long, striated and crested, ripening forests were not removed altogether. It is considered
from green to black. a relatively uncommon tree even in forest reserves
that have not or only lightly been logged. Regenera-
tion is probably dependent on episodal disturbance
Distribution
of the forest canopy and may seem poor over shorter
New Zealand: North Island (between Hokianga and time intervals.
Manukau Harbor and in the southern part of the IUCN: VU A1 (a, c, d)
Coromandel Peninsula).
TDWG codes: 51 NZN
Uses
Monoao is a podocarp with fine grained wood suit-
Ecology
able for construction timber, a use to which it was
Halocarpus kirkii is an uncommon forest tree occur- put in former times when it was still relatively abun-
ing in lowland subtropical mixed evergreen forest to dant around Auckland and in Northland. It is now
a maximum altitude of ca. 700 m. In the Waitakere protected from logging and most trees occur in for-
Ranges near Auckland the kauris (Agathis australis) est reserves. This species is not recorded in cultiva-
form emergent trees, with Dacrycarpus dacrydio- tion, but may be present in a few botanic gardens.
ides, Dacrydium cupressinum, Podocarpus cunning- According to Grimshaw & Bayton (2009: 388) there
hamii, Prumnopitys ferruginea, and ocasionally may be only one plant growing in the northern
Halocarpus kirkii together with various angiosperms hemisphere: at Tregrehan in Cornwall, England.
Juniperus L., Sp. Pl. 2: 1038. 1753. Type: Juniperus communis L. (Cupressaceae).

Sabina Mill., Gard. Dict., Abridg. Ed. 4, vol. 3. 1754.

Distribution
Type: Sabina vulgaris Antoine [Juniperus sabina
L.]; Arceuthos Antoine & Kotschy, Oesterr. Bot. Europe, Africa, Macaronesia, Asia, North America,
Wochen­bl. 4 (31): 249. 1854. Type: Arceuthos drupa- Central America, Caribbean.
cea (Labill.) Antoine & Kotschy [Juniperus drupacea
Labill.]; Sabinella Nakai, Chôsen Sanrin Kaihô 165:
Taxonomic notes
14. 1938. Type: Sabinella phoenicea (L.) Nakai [Juni-
perus phoenicea L.]. In my Monograph of Cupressaceae and Sciadopitys 381
(Farjon, 2005a), reference was made to the work of
Juniperus is the classical Latin name for junipers. R. P. Adams and collaborators, who have used bio-
chemical data, among other data, to delimit the spe-
cies in the genus Juniperus. I was critical of some
Description
of the species proposed on the basis of chemical
Decumbent or erect shrubs or trees, evergreen, compounds found in the terpines extracted from
monoecious or dioecious. Resin cavities in leaves. leaf samples. Since that critical review, the stream
Bark fissured, fibrous and stringy, exfoliating in of papers by Adams et al. proposing new taxa, or
long strips, or sometimes tesselated, hard and exfo- recombinations of existing names, has increased
liating in small flakes. Branches decumbent, erect, dramatically. DNA sequencing is now providing ad-
spreading or pendulous; foliage branches usually ir- ditional data, often interpreted in a way that result in
regularly disposed, sometimes forming more or less what are termed “cryptic” species: morphologically
flattened sprays. Leaves scale-like or acicular (the indistinguishable, but species nevertheless because
latter to 25 × 4 mm), decurrent or articulate, decus- the samples appear to represent distinct lineages in
sate or ternate (both leaf shapes and the two types of a phylogenetic (cladistic) analysis of the data. In the
phyllotaxis can occur on the same plant, but these context of this Handbook, it is impossible to enter
have either decurrent or articulate leaf bases), scale into a discourse about the various species concepts
leaves glandular or eglandular, leaf margins entire or scientists advocate or denounce (for an overview
minutely denticulate; apex obtuse or acute-pungent, see Stuessy, 2009 and for an entertaining debate on
amphistomatic or epistomatic. Pollen cones termi- this issue see Wheeler & Meier, 2000). However, it
nal or axillary, with terminal cones solitary and axil- seems to me safe to say that inferences about the de-
lary cones on dwarf shoots or sessile, 2–3 per ternate limitation of species when based on the ‘phylogeny’
acicular leaf whorl, subglobose to ovoid, 2–7 mm of DNA sequence data from (necessarily) limited
long; microsporophylls 6–18, decussate or ternate, sampling are at best controversial. It appears that we
with 2–6 abaxial pollen sacs. Seed cones terminal are confronted with the controversial enforcement
on axillary (dwarf) shoots, 4–30 mm, globose, semi- of phylogenetic monophyly (= holophyly) used here
globose or ovoid, indehiscent, hard dry or soft suc- to identify species. Are species merely genealogical
culent with a smooth or sometimes rugose, often lineages? Are species populations of interbreeding
glaucous or pruinose surface. Bract-scale complexes individuals, separate in that function from other
in 1–4 decussate or ternate whorls, entirely fused (in populations and thereby accumulating different
one species partly fused), often with visible sutures and eventually distinct characters? Are species just
and minute exserted bract apices on the surface. ‘kinds’, conjured up by us? “Cryptic” species sensu
Seeds 1–8 per cone, in one species fused to a single Adams et al. may seem to follow the first option with
‘stone’, often with resin vesicles and ridges but with- some evidence for accumulating genetic differences.
out wings. Cotyledons 2 (in a few species 3–6); juve- It is likely that they reflect ongoing evolution. Cryp-
nile leaves on seedlings or also on young to mature tic species certainly do present a problem in the
plants, together with scale leaves or exclusively. practicalities of how to recognize them. They are not
endorsed in this Handbook.
53 species.
 2a. Leaves articulate at middle. Seed cones (sub)
Synopsis
globose, with more than one seed (or seeds
The currently accepted classifications appear to agree fused to one ‘stone’) Group 1 (‘Oxycedrus’)
on a division of the genus in two sections: sect. Juni- 2b. Leaves decurrent at base. Seed cones ovoid-­
perus (syn. Oxycedrus) and sect. Sabina (but some- globose to ovoid, usually with a single seed
times three, with J. drupacea removed from sect. Group 2 (‘Squamata’)
Juniperus), but to differ in its further subdivisions
dependent on the character(s) chosen to separate
Key to the species of Group 1
these. In the Monograph of Cupressaceae and Sciado­
pitys (Farjon, 2005a), I presented some evidence in- 1a. Seed cones larger than 15 mm, hard, with well
382 dicating that these (sub)divisions may be ‘artificial’, marked scales in whorls of three; seeds fused
i.e. not based on or reflecting natural groups result- to a single ‘stone’. Pollen cones in clusters with
ing from evolution. None of the chracters used to whorls of small leaves. Habit a tree J. drupacea
separate these sections or subsections are consistent 1b. Seed cones smaller than 15 mm, or when larger
throughout and another choice of characters would (up to 23 mm), scales invisible or only marked
probably give another set of groups. A comprehen- as sutures; seeds free. Pollen cones solitary.
sive phylogenetic analysis of the genus is at this time Habit a shrub or tree 2
of writing still wanting (“in progress”, Adams et al., 2a. Leaves without a midrib on the adaxial side,
but see remarks above). A classification of the genus stomata in a single band 3
Juniperus should be based on inferred phylogeny 2b. Leaves with a midrib on the adaxial side, sepa-
as much as possible, but recognizing evolutionar- rating the stomata in two bands 4
ily significant character changes as well. We are still 3a. Stomatal band wider than the green margins of
some distance in time away from a robust phylogeny leaves J. communis
of distinct species which can inform a classification 3b. Stomatal band narrower than the green mar-
that is both informative and useful. gins of leaves, often hidden in a narrow groove
J. rigida
4a. Leaves usually obtuse. A species from Japanese
Key to the informal groups
islands in the Pacific J. taxifolia
The leaves referred to in the keys below (and with 4b. Leaves usually acute or pungent, if obtuse, very
few exceptions in the descriptions) are those found short (4–10 mm), boat-shaped 5
on (sub)ultimate foliage branchlets of mature plants. 5a. Branchlets pendulous (except in shrubs at high
Different shapes and sizes occur both on juvenile altitude). Habit usually a tree. A species from
plants and on so-called whip shoots, the faster grow- China (including Taiwan) J. formosana
ing leading shoots; leaves are generally larger on 5b. Branchlets spreading or sometimes pendulous.
older (lower order) branches, as they may continue Habit a shrub (rarely a tree) 6
to grow for a while with those branches. Plants in 6a. Leaves very short (4–10 mm), boat-shaped.
cultivation, even if true species, often tend to retain A species from the Azores J. brevifolia
juvenile leaf types longer, especially those grown 6b. Leaves at least 8 mm long, usually longer than
in a more humid climate than that of their natural 10 mm 7
range, or indeed in greenhouses. Such plants, espe- 7a. Leaves short or long (8–23 mm), widest near
cially when sterile, are often difficult to identify and the middle. A species from Madeira and the
these keys are not tuned to them, but to the species Canary Islands J. cedrus
as they are found in nature. 7b. Leaves mostly 10–20 mm long, widest near the
base. A species from the Mediterranean and the
1a. Leaves in mature plants all acicular (needle- Middle East J. oxycedrus
like, if sometimes short), in alternating whorls
of three 2
Key to the species of Group 2
1b. Leaves all scale-like, or at least some branchlets
with scale leaves in mature plants, decussate or 1a. Branchlets drooping or pendulous. A large
on some branchlets in whorls of three shrub or tree J. recurva
Group 3 (‘Sabina’)
1b. Branchlets spreading. A decumbent or erect 4a. Foliage branches often with scale leaves as well
shrub, occasionally a small tree 2 as acicular leaves, both types bearing male or
2a. Leaves small (2–5 mm), not more than five female cones J. przewalskii
times longer than wide, imbricate, usually ap- 4b. Foliage branches with scale leaves only
pressed; apex incurved J. pingii J. komarovii
2b. Leaves more than five times longer than wide, 5a. Ultimate branchlets subterete, rarely quadran-
spreading at a narrow angle to patent; apex gular 6
straight 3 5b. Ultimate branchlets mostly quadrangular (ex-
3a. Seeds usually 2–3 per cone, rarely a single seed cept branchlets with leaves in whorls of three).
J. procumbens A species of the Himalayas J. indica
3b. Invariably a single seed per cone J. squamata 6a. Habit usually a monopodial tree, a shrub only 383
under adverse ecological conditions. Leaves
sometimes in whorls of three. A species of the
Key to the subgroups of Group 3 and
Tibetan Plateau 51. J. tibetica
J. phoenicea
6b. Habit a decumbent or erect shrub, rarely a small
1a. Margins of scale leaves entire. Seed cones dark tree. Adult scale leaves all opposite-decussate.
blue-black, lustrous, succulent, containing a A species of Central Asia J. pseudosabina
single large seed (very rarely a second, smaller
seed) Group 3a
Key to the species of Group 3b (American)
1b. Character states not in this combination 2
2a. Margins of scale leaves minutely denticulate 1a. Ultimate branchlets usually > 1.5 mm thick;
(× 20!). Seed cones lighter coloured, often glau- leaves mostly in whorls of three, less often
cous, succulent or dry, containing 1–2 seeds ­opposite-decussate 2
(commonly only one seed) Group 3b 1b. Ultimate branchlets < 1.5 mm thick; leaves
2b. Margins of scale leaves entire. Seed cones < 10 mostly opposite-decussate, less often in whorls
mm diam., succulent, usually containing two of three 6
or more seeds (rarely only one) Group 3c 2a. Habit a shrub, rarely a tree 3
2c. Seed cones dry, containing more than two 2b. Habit a monopodial tree, rarely a shrub
seeds, often glaucous or pruinose, rarely succu- J. occidentalis
lent and red or purplish black 3 3a. Ultimate branchlets thick, 2-2.5 mm diam.,
3a. Margins of leaves minutely denticulate (× 20!) terete. Seed cones large, 10–18 mm J. californica
4 3b. Ultimate branchlets < 2 mm thick. Seed cones
3b. Margins of leaves entire Group 3e smaller, 4–13 mm 4
4a. Seed cones dry when fully mature, usually 4a. Seed cones dry, 7–13 mm. Habit sometimes a
glaucous Group 3d small, sturdy tree J. osteosperma
4b. Seed cones succulent, reddish or purplish black, 4b. Seed cones succulent when mature. Habit al-
containing 3–8 seeds J. phoenicea ways a shrub 5
5a. Seeds usually single in a cone. Shrub not coppic-
ing (sprouting from the base) J. monosperma
Key to the species of Group 3a (Asian)
5b. Seeds 1–2 per cone. Shrub readily coppicing
1a. Seed cones when mature < 10 mm long, glo- J. pinchotii
bose or ovoid 2 6a. Seed cones small, 4–7 mm, dry when mature 7
1b. Seed cones when mature usually > 10 mm long, 6b. Seed cones slightly larger, 5–9 mm, succulent
ovoid, rarely globose 3 when mature 8
2a. Ultimate branchlets slender, subterete, 0.8-1.2 7a. Bark becoming tesselated on lower stem of ma-
mm wide J. convallium ture small trees or large shrubs J. angosturana
2b. Ultimate branchlets coarser, quadrangular, 1.0- 7b. Bark becoming stringy, exfoliating in strips
1.7 mm wide J. saltuaria J. comitana
3a. Foliage branches drooping or pendulous 4 8a. Seed cones maturing to dark blue J. ashei
3b. Foliage branches spreading 5 8b. Seed cones maturing to pinkish or orange-red,
sometimes purplish blue 9
 9a. Glands of leaves small, rounded, flat and about 1b. Bark on the trunk of trees fibrous, exfoliating in
half as long as the decurrent leaf base long strips, soft 2
J. arizonica 2a. Habit a monopodial tree 3
9b. Glands of leaves large, rounded to oblong, flat 2b. Habit a shrub, rarely a small tree 4
or slightly raised and as long as the decurrent 3a. Foliage branches spreading. Seed cones 5-7.5
leaf base J. coahuilensis mm, brown (variously glaucous) J. jaliscana
3b. Foliage branches drooping to pendulous. Seed
cones 8.5-15 mm or larger, light brown (strong-
Key to the species of Group 3c (North
ly glaucous) J. flaccida
American, Caribbean, Eurasian)
4a. Seed cones light brown to reddish brown, dull,
384 1a. Foliage branchlets commonly with scale leaves with 1-3, rarely 4 seeds per cone 5
and/or acicular leaves on mature plants (some 4b. Seed cones usually purplish red or purplish
varieties of species have only one type of leaf) black, lustrous, with 2–9 seeds per cone
2 J. monticola
1b. Foliage branchlets of mature plants exclusively 5a. Leaves on ultimate branchlets small, nearly as
with scale leaves 3 wide as long, 0.8-1.2 × 0.8-1 mm, glands incon-
2a. Decumbent shrub. Foliage rarely with acicular spicuous J. durangensis
leaves (but more commonly in one part of the 5b. Leaves on ultimate branchlets larger, longer
range); seeds 1-3(-5) per cone J. sabina than wide, 1.2-1.5 × 1-1.2 mm, glands conspicu-
2b. (Decumbent) shrub or usually a large tree. Fo- ous but not with resin drops J. saltillensis
liage mostly with a mixture of scale leaves and
acicular leaves; seeds 2-4(-5) per cone
Key to the species of Group 3e (American,
J. chinensis
African, Eurasian)
3a. Seed cones often wider than tall, bilobed to
nearly reniform, with two diverging seeds 4 1a. All leaves on ultimate branchlets appressed,
3b. Seed cones all ovoid-globose to globose, with small and scale-like. Seed cones globose,
converging seeds J. virginiana 6–12(–14) mm. A tree of Eurasia J. excelsa
4a. Scale leaves on ultimate branchlets decussate or 1b. Characters not in this combination 2
in whorls of three, length/width ratio ≥ 3:1 2a. Habit a prostrate or decumbent shrub. A spe-
J. bermudiana cies of boreal North America J. horizontalis
4b. Scale leaves on ultimate branchlets all decus- 2b. Habit a (small or large) tree, rarely a shrub un-
sate, length/width ratio < 3:1 5 der adverse conditions 3
5a. Apices of scale leaves mostly mucronate or 3a. Seed cones usually broader than long, often
acute 6 more or less reniform or bilobed, in such cones
5b. Apices of scale leaves mostly obtuse 7 only two seeds present with diverging apices 4
6a. Apices of scale leaves mucronate or acute-acu- 3b. Seed cones globose or subglobose, with (2-)3-
minate. A species of the Caribbean J. gracilior 6(-8) seeds per cone with converging apices 5
6b. Apices of scale leaves acute or sometimes more 4a. Foliage with only acicular leaves (juvenile
or less obtuse. A species of W North America phase) but capable of bearing male or female
J. scopulorum cones. A species of the Caribbean J. saxicola
7a. Scale leaves more or less gibbous, gland often 4b. Foliage normally with scale leaves only
inconspicuous. A species of the Caribbean J. semiglobosa
J. barbadensis 5a. Scale leaves on foliage branchlets often with
7b. Scale leaves more flattened, gland always con- spreading, free apex J. foetidissima
spicuous. A species of Mexico J. blancoi 5b. Scale leaves on foliage branchlets all appressed
6
6a. Ultimate branchlets very slender, 0.6-1 mm
Key to the species of Group 3d (Mexican,
wide. Seed cones small, 3-7 mm diam
Arizona to W Texas)
J. procera
1a. Bark on (the lower part of) the trunk of trees 6b. Ultimate branchlets thicker, 1-1.4 mm wide.
tesselated or fissured, hard J. deppeana Seed cones usually larger, 6-10 mm diam 7
7a. Mature seed cones brown, often glaucous. mm long, terete; microsporophylls 10–12, decussate,
A species of Central America J. standleyi peltate; margins hyaline-denticulate; apex acumi-
7b. Mature seed cones blackish blue or purplish nate, bearing 3–4 abaxial pollen sacs. Seed cones
black, not glaucous J. thurifera numerous, terminal on very short, lateral branches
with only 6–8 decussate scale leaves, maturing in
one year, globose or subglobose to ovoid, 4–6 ×
Juniperus angosturana R. P. Adams, Biochem. 3–5 mm, purplish blue, then pinkish with glaucous
Syst. Ecol. 22 (7): 704. 1994. Juniperus monosperma bloom, drying brown. Bract-scale complexes 4(–6),
(Engelm.) Sarg. var. gracilis Martínez, Anales Inst. decussate, sometimes ternate, entirely fused with
Biol. Univ. Nac. México 17 (1): 111. 1946. Type: barely exserted bract tips, smooth (rugose in sicco),
Mexico: San Luis Potosí, Hacienda de Angostura, soft pulpy, slightly resinous. Seeds 1(–2), ovoid to 385
4 km W of Angostura, C. G. Pringle 3771 (holotype subglobose (when solitary), 3–5 × 2.5–4 mm, with
MEXU lost, isotype K). shallow grooves; apex more or less acute, lustrous
(light) brown, with a tan hilum at base.
Etymology
Distribution
The specific name refers to Hacienda de Angostura
in San Luis Potosí, Mexico, where the type specimen Mexico: Coahuila, Hidalgo, Nuevo León, San Luis
was collected. Potosí, Tamaulipas (Sierra Madre Oriental).
TDWG codes: 79 MXE-CO MXE-HI MXE-NL MXE-
SL MXE-TA
Vernacular names
No common names have been recorded for this spe-
Ecology
cies.
In open shrubland or in Pinyon-Juniper woodland,
associated with Pinus cembroides, Juniperus ­flaccida
Description
and sclerophyllous shrubs; on rocky slopes and
Small trees or shrubs to 3–8(–10) m tall, dioecious; along intermittent streams. Also found in open pine
trunk to 40–50 cm diam. Bark on trunk 1–2 cm forest with P. montezumae.
thick, fissured, usually tesselated, or with longer ex-
foliating plates; inner bark light brown or cinnamon;
Conservation
outer bark weathering grey. Branches thick, large,
spreading and assurgent, tortuous, foliage spread- IUCN: LC
ing or drooping, forming an irregular, spreading or
more or less dome-shaped crown. Foliage branch-
Uses
es iregularly disposed, lax and slender, ultimate
branchlets 4–20 mm long, straight or curved, 1–1.3 No uses have been recorded of this species.
mm wide, quadrangular to terete (depending on
phyllotaxis. Leaves decussate (mostly on ultimate
branchlets) or in alternating whorls of 3, imbricate, Juniperus arizonica (R. P. Adams) R. P. Adams,
appressed, or with a few reflexed leaf apices on older Phytologia 88 (3): 306. 2006. Juniperus coahuilensis
branchlets, oblong-rhombic or oblong-ovate on ul- (Martínez) Gaussen ex R. P. Adams var. arizonica
timate branchlets, 1–1.5 × 0.7 mm, obtuse or acute; R. P. Adams, Biochem. Syst. Ecol. 22 (7): 708. 1994.
upper margins minutely hyaline-serrulate (esp. of Type: USA: Arizona, Yavapai Co., Camp Verde,
smaller leaves); stomata few abaxially, near base, R. P. Adams 2132 (holotype BAYLU).
partially hidden by margins of lower leaves, adaxi-
ally in two bands from base to apex, separated by a
Etymology
narrow midrib, most leaves of ultimate branchlets
eglandular, some leaves with an oblong, inactive The species epithet refers to the State of Arizona,
gland, leaf colour yellowish green or light green. where the type specimen was collected.
Pollen cones terminal, solitary, ovoid-oblong, 2.5–3
 trees ­appear to rotrude distally from the cone =
Vernacular names
‘gymnocarpous’ cones).
No vernacular name has been given to this species;
an obvious choice would be Arizona juniper.
Taxonomic notes
This taxon, formerly recognized as a variety under
Description
Juniperus coahuilensis (see e.g. Adams, 2004, Far-
Large shrub to 5 m, or small tree to 10 m, dioecious, jon, 2005a), has been found in a recent phylogenetic
rarely monoecious; multistemmed or with a short analysis based on DNA sequence data of one-seed-
trunk and a maximal diam. of 50 cm. Bark of thick ed, serrate-leaf junipers of the SW USA and north-
386 branches and trunk fibrous, exfoliating in strips, ern Mexico to be quite unrelated to that species. It
weathering ash-grey. Branches numerous, long, as- appears to be forming a clade with J. occidentalis
cending from base or spreading, often curved, those and J. osteosperma, whereas J. coahuilensis is nearer
of higher orders ascending or spreading, forming a J. monosperma and J. pinchotii. Therefore, Adams
broad, rounded or more irregular and open crown. elevated it to species rank, which on the given evi-
Foliage branches numerous, irregularly disposed dence seems to be convincing and is here followed.
but not pendulous, ultimate branchlets spreading Its morphology is, however, most similar to that of
30–70 degrees, slender and stiff, up to 15 mm long, J. coahuilensis and the two may not be so easy to
1–1.5 mm thick, (weakly) quadrangular in cross sec- separate in the field, as the only reported consistent
tion, covered with scale leaves, persistent. Leaves difference is in the size of the leaf glands.
decussate on all ultimate branchlets, decussate or
in alternate whorls of 3 on older branchlets, imbri-
Distribution
cate, scale-like, 1.2–1.7(–1.8) × 0.7–1.2 mm on lateral
branchlets, decurrent, rhombic or ovoid-rhombic; SW USA: Arizona, SW New Mexico, W Texas.
apex appressed or rarely free, obtuse to acute; mar- TDWG codes: 76 ARI 77 NWM TEX
gins minutely denticulate; stomata on abaxial side
limited to two small spots near leaf base, on adax-
Ecology
ial surface in two tapering bands; glands central on
small scale leaves, below the free apex on longer In Bouteloua grassland and on adjacent rocky slopes.
whip shoot leaves, more or less conspicuous, small, It has a capacity to coppice, considered ‘weedy’ in
rounded, flat, half as long as the decurrent part of some areas by sheepmen and cattlemen. In Arizona it
the leaf; exudate sometimes present; leaf colour yel- occurs with Juniperus osteosperma in some localities;
lowish green or light green. Pollen cones numerous, other associated species are sometimes Opuntia spp.
terminal, solitary, ellipsoid or ovoid-oblong, 3–4 × or Yucca spp., but commonly its only associates are
2–2.5 mm, yellowish maturing to light brown; mi- grasses. Altitudinal range 980–1600(–2200) m a.s.l.
crosporophylls 8–10, decussate, peltate, acute, with
minutely denticulate upper margins and with 3–4 Conservation
abaxial pollen sacs. Seed cones terminal on straight,
IUCN: LC
very short branchlets, maturing in one season at-
taining variable colours from pinkish(–red) or or-
Uses
ange(–red) to purplish blue, usually glaucous, (sub-)
globose or sometimes broad ovoid, 5–7 mm diam. No uses have been recorded of this species.
(if ovoid to 9 mm long), internally soft pulpy, suc-
culent, yellowish to orange-brown. Bract-scale
complexes 4(–6), entirely fused, decussate, bract Juniperus ashei J. T. Buchholz, Bot. Gaz.
apices minutely exserted or hidden; surface smooth, (Crawfordsville) 90 (3): 329. 1930.
rugose when dry. Seeds 1(–2) per cone, ovoid-glo-
bose, 3–5(–6) × 3–4(–5) mm, base rounded; apex
Etymology
acutish, shallowly grooved and ridged on sides, not
or slightly resinous, chestnut brown, with a large, The species epithet commemorates William W. Ashe,
lighter lobed hilum proximally (the seeds in some who made the first botanical collections of it.
 Vernacular names Ecology
Ashe juniper, Mountain cedar, Ozark white cedar, This species occurs in open grassland and shrub-
Rock cedar, Post cedar, Mexican juniper land, sometimes associated with Juniperus pinchotii,
Pinus remota, and Quercus spp.; in sandy or rocky
soil on plains, on bluffs and ridges, on hillsides or
Description
along stream beds in sandy or rocky soil, commonly
Large shrubs or small trees 6–10(–15) m, dioe- over limestone. The altitudinal range is (150–)600–
cious; with a short trunk 1–3 m tall and a diam. 1550 m a.s.l. The climate is continental, with warm
of 30–50(–120) cm, or branching at base. Bark of summers and cold winters, drier with hotter sum-
of thick branches and trunk exfoliating in strips, mers in the south of its range. 387
brown weathering grey. Branches long, spreading
to ascending or nearly erect, forming a sympodial,
Uses
broad, rounded and dense or more irregular and
open crown. Foliage branches numerous, irregularly In the past the wood was used by cattle ranchers for
disposed but not pendulous, ultimate branchlets fenceposts, but metal has displaced it. The wood is
spreading to erect, stiff, 5–10(–20) × 0.9–1.3 mm, now mainly used to produce ‘cedarwood oil’ through
mostly 4-sided in cross section, covered with closely steam distillation. This species is not known to be in
appressed leaves, persistent. Leaves decussate, or oc- cultivation, but it may be present in a few botanic
casionally in alternate whorls of 3 on whip shoots, gardens.
(slightly) imbricate, scale-like, 1–2 × 0.8–1.2 mm on
lateral branchlets, decurrent, rhombic, often keeled, 2 varieties are recognized:
acute; margins minutely denticulate; stomata on the
abaxial side limited to decurrent leaf base, on the Juniperus ashei J. T. Buchholz var. ashei. Sabina
adaxial surface in two bands; glands obscure or con- ashei (J. T. Buchholz) A. V. Bobrov & Melikyan,
spicuous on whip shoot leaves, hemispheric or oval Komarovia 4: 81. 2006. Type not designated.
to elongate, raised; exudate absent; leaf colour light
to dark green. Pollen cones numerous, terminal,
Description
solitary, subglobose to ovoid, 2–4 × 2 mm; micro-
sporophylls 6–10, decussate, peltate with minutely Glands on whip shoots hemispheric. Seed cones 7–9
erose-denticulate or entire margins and with 3–4 mm diam.; seeds mostly 1 per cone, rarely 2.
(–5) abaxial, large pollen sacs. Seed cones terminal
on straight short branchlets, maturing to dark blue
Distribution
within 1 year, globose to broadly ovoid, (5–)6–9 mm,
succulent and resinous. Bract-scale complexes en- Central USA: Arkansas, Missouri (Ozark Mts.),
tirely fused, 4–6, decussate, indiscernible in mature Oklahoma, Texas (Edwards Plateau).
cones; surface smooth, rugose when dry, lustrous TDWG codes: 74 MSO OKL 77 TEX 78 ARK
or very glaucous (in some populations brownish).
Seeds 1(–2, rarely 3) per cone, broad ovoid, 4–6 × Conservation
3–4.5 mm, not flattened, smooth or vaguely grooved,
IUCN: LC
lustrous yellowish brown to chestnut brown, with a
lighter hilum proximally.
Juniperus ashei J. T. Buchholz var. ovata
R. P. Adams, Phytologia 89 (1): 17. 2007. Type: USA:
Distribution Texas, Crockett Co., 5 km W of Ozona, R. P. Adams
7463 (holotype BAYLU).
Central USA: Arkansas, Missouri (Ozark Mts.),
Oklahoma, Texas (Edwards Plateau, W Texas); Mex-
Description
ico: N Coahuila.
TDWG codes: 74 MSO OKL 77 TEX 78 ARK 79 Glands on whip shoots oval to elongate. Seed cones
MXE-CO (5–)6(–8) mm diam.; seeds more often 2 instead of
1 per cone.
 ­ ecussate, entirely fused, sutures or bract tips invis-
d
Distribution
ible. Seeds (1–)2–3(–4) per cone, ovoid-globose (if
USA: W Texas; Mexico: extreme N Coahuila. more than two in a cone flattened on one side), 2–3
TDWG codes: 77 TEX 79 MXE-CO mm, light yellowish brown.

Conservation Distribution
IUCN: NE Bahamas, Cuba, Haiti, Jamaica, Windward Islands.
TDWG codes: 81 BAH CUB HAI-HA JAM WIN-SL

388 Juniperus barbadensis L., Sp. Pl. 2: 1039. 1753.

Ecology
In shrubland and in ‘pine barrens’ or pine savan-
Etymology
nas, in the latter associated with Pinus caribaea var.
The species epithet refers to the island of Barbados. ­bahamemsis, P. tropicalis, in upland hills with P. car-
ibaea var. caribaea or P. occidentalis, also scattered
on rocky cliffs and escarpments in nutrient poor
Vernacular names
sand or rocks, commonly over limestone or karst. In
West Indies juniper; Barbados cedar (var. barbaden- the pine savannas fires are very frequent. Its altitu-
sis), Red cedar; Cèdre (French) (var. lucayana) dinal range is from 30 m to 1530 m a.s.l. The climate
is tropical, with a mean annual temperature above
20 °C and annual precipitation ca. 1000–1800 mm
Description
with a prolonged dry season.
Shrubs or trees to 10–12 m tall; trunk to 40 cm
diam., dioecious. Bark on tree trunks thin, stringy, 2 varieties are recognized:
exfoliating in long strips or thin plates, becoming
grey-brown. Branches spreading horizontally or Juniperus barbadensis L. var. barbadensis. Type:
ascending, in shrubs ascending or erect, forming Barbados: locality unknown, [if collected from
initially pyramidal, then open and irregular crowns Barbados, it is extinct there], leg. ign. LINN 1198.1
in trees. Foliage branchlets numerous, irregularly (lectotype LINN).
disposed or sometimes more or less distichous and
gradually shorter, variable in length from 5–30(–60)
Description
mm, slender or very slender, 0.8–1.2 mm diam., sub-
terete to quadrangular in branchlets with scale leaves, Ultimate branchlets slender, 1.0–1.2 mm diam.
persistent. Leaves decussate, decurrent, scale-like on Leaves imbricate, slightly spreading, forming 4 reg-
mature plants and branches, imbricate, appressed ular rows of nearly free leaf tips along the branch-
but slightly diverging, on ultimate branchlets 1–1.5 lets; apex obtuse or acute; glands inconspicuous or
mm long, rhombic to rhombic-lanceolate; margins conspicuous.
entire; apex obtuse to acute or mucronate, abaxially
with a more or less conspicuous rounded to elliptic
Distribution
gland, green or greyish green; stomata few at base
abaxially, in one or two concave fields adaxially. Pol- Windward Islands: St. Lucia (Petit Piton); extinct on
len cones terminal, oblong, 2.5–3 × 1.5 mm, more or Barbados.
less quadrangular; microsporophylls 12–16, decus- TDWG codes: 81 WIN-SL
sate, broad peltate, with rounded and entire upper
margin, abaxially bearing 2–3 globose pollen sacs.
Ecology
Seed cones terminal on erect, short ultimate branch-
lets, maturing in one season, becoming irregularly In its only remaining habitat restricted to limestone
globose or broad pyriform to nearly reniform, 4–5 cliffs near the summit of a mountain, with other
× 6–7(–8) mm, usually soft pulpy, resinous, pru- shrubs.
inose-blue or blackish blue. Bract-scale complexes 4,
 Conservation Uses
The population that apparently once existed on Bar- The wood is locally used for the carving of souve-
bados disappeared probably in the early part of the nirs.
18th century, the location having been converted to
sugar cane plantations. Recently this variety has
been found on the ‘Petit Piton’ summit on St. Lucia Juniperus bermudiana L., Sp. Pl. 2: 1039. 1753. Type:
at ca. 730 m a.s.l., where Adams (1989) found ca. 25 Netherlands: (cultivated), leg. ign. [ex herb. D. van
trees, all within 30 m of the rocky summit, where Royen] 901.130-394 (neotype L).
they seem to have escaped attention from goats as
well as people. Despite the possibility of its occur- 389
Etymology
rence on other nearby islands, nothing has been
found there to date, and this appears to be the only The species epithet refers to its origin, the island of
extant population of this variety. Bermuda, where it is endemic.
IUCN: CR (D)
Vernacular names
Juniperus barbadensis L. var. lucayana (Britton)
R. P. Adams, Phytologia 78: 145. 1995. Juniperus Bermuda cedar, Bermuda red cedar, Bermuda juni-
lucayana Britton, N. Amer. Trees: 121. 1908. per
Type: Bahamas: New Providence, Southwest Bay,
N. L. Britton & L. J. K. Brace 497 (holotype NY).
Description
Trees to 10–12 m tall, with monopodial, erect trunk
Description
to 60 cm diam., dioecious. Bark on tree trunks thin,
Ultimate branchlets very slender, 0.8–1.0 mm diam. exfoliating in strips, red-brown becoming grey-
Leaves imbricate, appressed or slightly spreading, brown. Branches spreading horizontally or ascend-
less markedly in 4 rows; apex obtuse to acute or ing, long, forming initially pyramidal, then open and
mucronate; glands conspicuous. irregular, spreading or flat-topped crowns. Foliage
branchlets numerous, disposed at angles of 25–35
degrees, ultimate branchlets variable in length from
Distribution
10–40 mm, lax, 1.3–1.6 mm diam., quadrangular
Bahamas, Cuba (Sierra de Nipe, Buenos Aires, Isla or sometimes subterete, persistent. Leaves on ulti-
de la Juventud [Isla de Pinos]), Haiti, Jamaica. mate branchlets decussate, occasionally in alternat-
TDWG codes: 81 BAH CUB HAI-HA JAM ing whorls of 3, decurrent, imbricate, appressed but
slightly diverging, overlapping a third to half of the
next leaves, on ultimate branchlets scale-like, 1.5–2.5
Conservation
× 1–1.5 mm, ovate-rhombic to rhombic-­lanceolate;
This, the more widespread form of J. barbadensis, is margins entire; apex obtuse to acute-mucronate;
nevertheless severely restricted in most of its range leaves abaxially with a conspicuous elliptic to lin-
and threatened by habitat changes. Small stands of ear gland, green; stomata few at base abaxially, in
5–30 trees are the usual situation in the Bahamas two concave fields adaxially. Pollen cones terminal
and in Jamaica (Adams, 1989) and some are also on ultimate branchlets, oblong-cylindrical, 4–6 ×
threatened by cutting as the wood is used in sou- 2–3 mm, more or less quadrangular; microspo-
venir carving. In Cuba, larger populations exist in rophylls 12–16, decussate, peltate; upper margin
the Sierra de Nipe and in swampy areas on Isla de la rounded and entire-hyaline, abaxially bearing 4–6
Juventud [Isla de Pinos], but an accurate assessment flattened, tightly packed pollen sacs. Seed cones
of this taxon in Cuba has not been undertaken to terminal on erect, short ultimate branchlets, matur-
date. At least some stands in Cuba are in less acces- ing in one season, becoming irregularly globose or
sible locations, providing for the time being some broad pyri­form to nearly reniform, 4–6 × 5–8 mm,
measure of protection. usually soft pulpy, resinous, pruinose-blue or dark
IUCN: VU (B1+2c) purplish blue. ­Bract-scale complexes 6, decussate,
 rarely in 2 whorls, entirely fused, sutures invisible, a Juniperus blancoi Martínez, Anales Inst. Biol. Univ.
few minute bract tips protruding. Seeds 1–2(–3) per Nac. México 17 (1): 73. 1946.
cone, ovoid-globose, 2–3 mm, more or less keeled,
lustrous brown.
Etymology
This species was named after Cenobio E. Blanco,
Distribution
who collected the type specimen.
Bermuda.
TDWG codes: 81 BER
Vernacular names
390 No common names were recorded for this species.
Ecology
Relict trees occur on eroded limestone rocks with
Description
pockets or fissures filled with sandy soil, or on hills
with skeletal, sandy soil, from near sea level to 50 m Small (shrubby) trees 8–10(–15) m, or sometimes a
a.s.l. small shrub, dioecious; (short) trunk with a maximal
diam. of 50 cm. Bark on larger trunks longitudinally
fissured, rough, exfoliating in thin, elongated strips
Conservation
or more compact, brown weathering grey-brown.
In the late 1940’s populations became heavily infest- Branches spreading, or ascending in young trees, of-
ed with two accidentally introduced species of ‘scale’ ten contorted, forming a pyramidal crown in young
insects, of which the Juniper Scale (Carulapsis mini- trees, to a broad, rounded or more irregular crown
ma) outcompeted the other species and became le- in old trees. Foliage branches numerous, irregu-
thal to 99% of the then existing junipers. Introduced larly disposed, spreading to subpendulous, ultimate
and naturalized trees and shrubs (Citharexylum branchlets spreading or drooping, lax, slender, 4–25
spinosum, Eugenia uniflora, Pimenta officinalis, and mm long, 0.8–1.2 mm thick, more or less quadran-
Schinus terebinthefolius) have largely replaced the gular in cross section, covered with scale leaves, per-
died stands of Bermuda cedar. Some trees survived, sistent. Leaves decussate, imbricate, scale-like, 1.3–2
possibly due to some resistance to the insects, but × 0.7–1 mm on lateral branchlets, decurrent, oblong-
regeneration is largely unsuccessful. Where regen- rhombic; apex appressed or just free, obtuse or api-
eration has occurred, young trees may have devel- culate; margins (hyaline-)entire; stomata on abaxial
oped some resistance but this needs confirmation. side limited to leaf base, on adaxial surface in two
Introduction of pest-resistant Juniperus virginiana tapering bands; glands conspicuous, large, elliptical
could pose a threat to the genetic integrity of still or oblong, slightly depressed, usually inactive; leaf
existing trees of J. bermudiana. Ongoing urbanisa- colour yellowish green to grey-green. Pollen cones
tion has left little natural habitat on the island but numerous, terminal, solitary, ovoid or subglobose,
some nature reserves have been established. 2–3 × 1.5–2 mm; microsporophylls 6–8, decussate,
IUCN: CR (B1+2c) peltate, with entire or hyaline-erose upper margins
and obtuse or mucronate apex; 3–4 abaxial pollen
sacs. Seed cones terminal on straight or curved, 3–6
Uses
mm long branchlets, maturing to purplish red, dark
When Bermuda cedar was still abundant the red- blue or tan with glaucous bloom in 1–2 years, sub-
dish, durable and sometimes beautifully figured globose to reniform or bilobed, 5–7 × 5–9 mm (often
wood was used in shipbuilding and especially cabi- wider than long), internally soft pulpy. Bract-scale
net making. Cabinets made of its wood are highly complexes 4–6, entirely fused, decussate, mostly in-
valued and treated as heirlooms in Bermuda. This discernible in mature cones; bract apices minutely
species has been taken into cultivation from the late exserted; surface smooth, rugose when dry, usually
17th century but only survives in countries with very glaucous. Seeds 1–2(–5) per cone, with apices often
mild winters, such as southern Europe, Ireland, Cal- diverging, irregularly ovoid-triangular or curved at
ifornia and Florida. apex, sometimes concavo-convex, 3–5 × 2–4 mm,
grooved and ridged on sides, with small resin pits Juniperus blancoi Martínez var. huehuentensis
near base, light brown, with a large darker hilum R. P. Adams, Biochem. Syst. Ecol. 34: 207. 2006.
proximally. Type: Mexico: Durango, Cerro Huehuento, summit
S of Huachichiles, S. Gonzáles et al. 6832 (holotype
CIIDIR).
Distribution
Mexico: Chihuahua, Sonora, Durango, México.
Description
TDWG codes: 79 MXC-ME MXE-CU MXE-DU MXN-
SO Shrubs 0.5–1.5 m tall, 2–4 m wide. Most seed cones
situated on the underside of branches.
391
Ecology
Distribution
The ecology of this species has not been assessed
due to a paucity of specimen data. It is believed to Mexico, Durango (Sierra Madre Occidental, sum-
occur in the drier types of pine woodland with Pinus mit area of Cerro Huehuento).
cembroides. TDWG codes: 79 MXE-DU

Uses Ecology
No uses have been recorded of this species. This is a subalpine shrubby form of J. blancoi pres-
ently only known from one mountain summit area
3 varieties are recognized: at 3150–3270 m a.s.l., where it grows together with
J. deppeana var. robusta in nearly bare rock and
Juniperus blancoi Martínez var. blancoi. Type: scree.
Mexico: Durango, El Salto Mun., El Salto, Arroyo
de Peñuelas, M. Martínez & C. E. Blanco 10527
Conservation
(holotype MEXU).
IUCN: NE
Description
Juniperus blancoi Martínez var. mucronata
Small trees. Leaves on ultimate branchlets oblong- (R. P. Adams) Farjon, Monogr. Cupressaceae
rhombic or rhombic, obtuse, appressed. Mature & Sciadopitys: 249. 2005. Juniperus mucronata
seed cones tan or blue, with glaucous bloom. R. P. Adams, Biochem. Syst. Ecol. 28 (1): 158.
2000. Type: Mexico: Sonora, Maicoba River, 19
km W of Maicoba, along Mex. Hwy. 16 at km 307,
Distribution
R. P. Adams 8704 (holotype SRCG).
Mexico: Durango (El Salto), México (Carmona).
TDWG codes: 79 MXC-ME MXE-DU
Description
Small trees. Leaves on ultimate branchlets rhombic
Conservation
or ovoid-rhombic, apiculate; apex just free. Ma-
This taxon is only known from less than five loca- ture seed cones purplish red or blue, with glaucous
tions, but in some of these it is locally common and bloom.
no particular threats seem to be imposed on it.
IUCN: VU (D2)
Distribution
Mexico: Chihuahua, Sonora.
TDWG codes: 79 MXE-CU MXN-SO
 10–14, in alternating whorls of 3, peltate, with erose-
Conservation
denticulate margins and acute apex, bearing 3–4
This taxon is only known from two collections made abaxial pollen sacs near lower margin. Seed cones
by Robert P. Adams, on 20 December 1998 only a maturing in 2 years; mature cones globose, 6–9 mm
few km separate from each other. It is likely that diam., with 1(–2) whorls of 3 completely fused bract-
this variety occurs more abundantly in this area of scale complexes, only the upper whorl fully devel-
northern Mexico, but a search of the herbaria has oped, 3 sutures visible on distal part of cone, bract
not resulted in further specimens. Therefore its ex- tip hidden or very small, tissue soft pulpy, juicy in
tent of occurrence (EOO) is likely to be small, per- most cones; immature cones green, hard; mature
haps less than 100 km2, and its area of occupancy cones dark red-brown, often slightly glaucous. Seeds
392 (AOO) smaller than 20 km2 with fewer than 1000 1–3 per cone, (angular-)ovoid, 5–6 × 3–4 mm, light
mature individuals. brown with obtuse apex.
IUCN: VU (D1, 2)
Distribution
Juniperus brevifolia (Seub.) Antoine, Cupress.- Azores.
Gatt.: 16, t. 20–22. 1857. Type: Macaronesia: Azores, TDWG codes: 21 AZO
Flores, [the specimen at P is recorded as from
Faial], C. F. Hochstetter 124 (holotype not located,
Ecology
isotypes K, P).
The habitat of this species has apparently been al-
tered greatly since the arrival of man and his domes-
Etymology
tic animals on the islands in the 16th century. It is a
The species epithet means “with short leaves”. constituent of the Macaronesian evergreen forest of
which only scattered remnants remain. Important
species in this forest type in the Azores are Laurus
Vernacular names
azorica, Persea indica, Myrica faya, Erica azorica,
Azores juniper, Cedro (Portugese) Frangula azorica, Ilex perado subsp. azorica, and
shrubs such as Bencomia, Pericallis and Phyllis. Well
developed forest occurs often on newly formed lava
Description
flows. The altitudinal range of J. brevifolia is 240–
Spreading or tall shrubs to small trees, to 6 m, dio- 800 m a.s.l. The prevailing SW winds (‘westerlies’)
ecious; trunk diam. to 50 cm, low branching. Bark set the conditions for a moist but mild climate with
on large trunks with long, soft strips, from reddish abundant rain and fog.
brown to grey-brown. Branches numerous, spread-
ing, assurgent or ascending. Foliage dense and stiff,
Conservation
with numerous spreading, ‘jointed’ ultimate branch-
lets, persistent, forming a shrubby, irregular crown Forest destruction and fragmentation of remnants
(prostrate at high altitudes). Leaves in alternating of natural vegetation have all but eliminated large
whorls of 3, rarely 4, non-decurrent, spreading 45–90 trees of Juniperus brevifolia. Forest remnants are
degrees at nodes 1.5–3(–5) mm apart, rigid, jointed now very restricted and scattered. Afforestation
to leaf part adnate with shoot, lanceolate to narrowly with exotic trees, e.g. Cryptomeria and Eucalyptus, is
ovate (‘boat-shaped’), 4–10 × 1–3 mm, usually broad- widespread and causes (irreversible?) habitat chang-
est near the curved, slightly thickened base; margins es unsuitable to native trees. Invasive shrubs, e.g.
entire; abaxial face keeled; leaf colour green to dark ­Hydrangea macro­phylla, spread over semi-­natural
green; apex obtuse-acuminate or acute-pungent; vegetation and ubiquitous grazing of domestic ani-
epistomatic, with stomata in two conspicuous bands mals cause further habitat degradation. Today Ju-
with whitish cuticular wax separated by a midrib, niperus brevifolia is only known as a (straggling)
bordered by green margins broader than the midrib. shrub, but remains of old trunks on some islands
Pollen cones solitary, 1–2 per leaf whorl, subglobose indicate its former extent and condition.
to ovoid-oblong, 3–5 × 2–4 mm; microsporophylls IUCN: EN (B1+2c)
 18), in alternating whorls of 3, peltate, with minutely
Uses
erose-denticulate or entire margins and with 3–5
This shrubby juniper is rare in cultivation; it would abaxial pollen sacs. Seed cones often numerous on
be suitable for gardens in areas with mild winters. whole plant, terminal on straight short branchlets,
Its endangered status in the Azores merits wider use maturing to purplish brown with whitish pruinose
in cultivation especially in connection with ex situ bloom in second season, globose to ovoid, 10–18 ×
conservation programmes. 7–11 mm, relatively dry-fibrous, slightly resinous.
Bract-scale complexes entirely fused, 6, in 2 alternat-
ing whorls of 3, partly discernible in mature cones,
Juniperus californica Carrière, Rev. Hort., sér. 4, 3: bract apices exserted, subterminal, causing slight ris-
352. 1854. Type: Illustration in Rev. Hort., sér. 4, 3: es in scale tissue, surface smooth, rugose when dry, 393
353, f. 21. 1854 (lectotype). Fig. 132, 133 often very glaucous. Seeds 1(–2, rarely 3) per cone,
broadly triangular-ovoid, 5–8(–9) × 4–7 mm, with
Juniperus californica Carrière f. lutheyana J. T. How- flattened base and acutish apex, grooved or pitted on
ell & Twisselm., Four Seasons 2 (4): 16. 1968. sides, lustrous yellowish brown to chestnut brown,
with a large, lighter 3-lobed hilum proximally.
Etymology
Distribution
The species epithet denotes its origin, the US State
of California. USA: W Arizona, California, S Nevada; Mexico:
Baja California Norte (including Cedros Island and
Guadalupe Island).
Vernacular names
TDWG codes: 76 ARI CAL NEV 79 MXI-GU MXN-
California Juniper; Huata, Cedro (Spanish) BC

Description Ecology
Large shrubs to 5–7(–10) m, dioecious, sometimes This species is adapted to one of the driest habitats
monoecious; with a short trunk to 1 m tall and a in which Juniperus can still survive well. It is locally
diam. of 30–40 cm, but more often multistemmed. common in desert scrubland of the Colorado, Mo-
Bark of thick branches and trunk exfoliating in have and Sonoran Deserts (‘Upper Sonoran Desert
strips, weathering grey. Branches numerous, long, scrub’) but extends into chaparral and open Pinus
ascending or nearly erect, forming a broad, rounded attenuata or P. sabiniana woodland in somewhat
and dense or more irregular and open crown. Fo- more mesic sites, which often occur nearer the Pacif-
liage branches numerous, irregularly disposed but ic coast. The altitudinal range is 500–1400 m a.s.l. In
not pendulous, ultimate branchlets spreading to (semi-)desert vegetation its common associates are
erect, very stout and stiff, up to 40 mm long, 2–2.5 Yucca brevifolia, Y. schidigera, Agave deserti, and Op-
mm thick, subterete or rarely weakly 4-sided in cross untia spp., with Larrea divaricata subsp. tri­dentata in
section, covered with closely appressed leaves, per- the lower, hotter basins and Seriphidium tridentatum
sistent. Leaves in alternate whorls of 3, rarely oppo- (Artimisia tridentata) and Chryso­thamnus nauseo-
site-decussate, imbricate, scale-like, 1.5–3 × 1.5–2.5 sus in cooler uplands. In Pine-Juniper woodlands
mm on lateral branchlets, decurrent, rhombic to Quercus spp., Ceanothus leucodermis and Arcto­
ovoid-rhombic, gibbous, acutish; margins minutely staphylos glauca are commonly growing with it. This
denticulate; stomata on the abaxial side limited to de- juniper can occur on barren serpentine or among
current leaf base, on adaxial surface in two tapering granite boulders but is not found in rock crevices, as
bands; glands conspicuous, oval or oblong, slightly it needs (coarse) alluvial material to spread its roots.
depressed, exudate sometimes present; leaf colour In much of its range there is a long, dry summer pe-
yellowish green or light green. Pollen cones numer- riod and rains occur only in winter, in some interior
ous, terminal, solitary, sub­globose to ovoid-oblong, desert valleys rain is erratic and the junipers may be
3.5–6.5 × 2–3 mm; microsporophylls (10–)12–15(– associated with deeper water sources in alluvial fans.
 (weakly) keeled or grooved, green to dark green;
Conservation
apex acuminate or acute-pungent; epistomatic, sto-
A widespread species not in danger of extinction. It mata in two conspicuous bands separated by a mid-
is possibly extinct on Guadalupe Island. rib, bordered by green margins broader than midrib.
IUCN: LC Pollen cones solitary, 1–2 per leaf whorl, subglobose
to ovoid-oblong, 3–6 × 2–5 mm, orange-brown;
microsporophylls 10–14, in alternating whorls of 3,
Uses
peltate, with erose-denticulate margins, bearing 3–4
No commercial uses have been recorded; it could abaxial pollen sacs near lower margin. Seed cones
make an attractive ornamental shrub in hot, dry cli- axillary on short (1–2 mm) dwarf shoots with whorls
394 mate and is occasionally used in garden landscaping of minute scale leaves, maturing in 2 years; mature
and for bonsai training in California, USA. In the cones globose, 8–14 mm diam., with 1(–2) whorls of
past, the wood was used for fence posts, but these 3 completely fused bract-scale complexes and only
are now all of metal. the upper whorl fully developed, 3 sutures visible on
distal part of cone; bract tip hidden or very small;
tissue soft pulpy, more or less resinous; immature
Juniperus cedrus Webb & Berthel., Hist. Nat. Iles cones green, hard, mature cones orange-brown.
Canaries 3 (2), Phytogr. Canar. 3 (89): 277. 1847. Seeds 1–3 per cone, ± ovoid, 5–10 × 4–6 mm.
Type: Macaronesia: Canary Islands, [“Ins. Can.”],
P. B. Webb s.n. (holotype not located, isotype K).
Taxonomic notes
Juniperus oxycedrus L. subsp. maderensis Menezes, The variation found in the size and shape of the
Bull. Acad. Int. Geogr. Bot. 18: xii. 1908; Juniperus ce- leaves of this species is considerable, from narrow
drus Webb & Berthel. subsp. maderensis (Menezes) (ca. 1 mm) and long (23 mm), pungent to broad (2.2
Rivas Mart. et al., Itinera Geobot. 15 (2): 703. 2002. mm) and short (8 mm), obtuse. As in J. oxycedrus, to
which it is somewhat more distantly related, length
and acuteness of the leaves generally decrease with
Etymology
increasing age of the trees and probably with in-
The species epithet is the classical name for the wood creasing exposure to light and weather. Specimens
of cypresses and/or junipers. seen from Madeira have usually short leaves, but
these are straight, not curved, as is common in spec-
imens from the Azores (J. brevifolia).
Vernacular names
Canary Islands juniper
Distribution
Canary Islands (Tenerife, La Palma, Gran Canaria,
Description
Gomera), Madeira.
Trees to 20–25 m tall (most surviving plants smaller TDWG codes: 21 CNY MDR
and shrubby), dioecious; usually monopodial, often
low branching. Bark on trunks with long, soft strips,
Ecology
grey-brown. Branches long, spreading, assurgent or
ascending. Foliage usually sparse, with spreading or Originally this species would have been a constitu-
pendulous, ‘jointed’ ultimate branchlets, persistent, ent of the Macaronesian evergreen forest, but as this
forming an open, irregular and broad crown (young vegetation type, and particularly the more open,
trees more pyramidal). Leaves in alternating whorls drier variant in which Juniperus cedrus had its op-
of 3, rarely 4, non-decurrent, spreading 30–60(–90) timum, has been altered or has disappeared alto-
degrees at nodes 3–8(–10) mm apart, rigid, jointed gether, it is now confined to steep rocky places at
to leaf part adnate with greenish yellow or glaucous altitudes ranging from 450 m to 2400 m a.s.l. Here
shoot, linear-lanceolate to narrowly ovate (‘boat- regeneration can succeed, but a forest with junipers
shaped’), (4–)8–23 × 1–2.3 mm, straight, usually and other trees will not easily develop.
broadest near middle; margins entire; abaxial face
 then open and irregular crowns in trees. Foliage
Conservation
branchlets numerous, spreading, assurgent or erect
This species is now restricted to inaccessible places in shrubs, spreading to pendulous in trees, slen-
both in the Canary Islands and Madeira. According der, 1–1.5 mm diam., subterete to quadrangular in
to David Bramwell, Director of the Botanic Garden branchlets with scale leaves, persistent. Leaves dec-
in Las Palmas, there are probably not more than 100 ussate or in alternating whorls of 3, short decurrent,
mature trees remaining in the wild (on Gran Canaria of two types: needle-like and scale-like. Needle-like
less than 6, numbers on Las Palmas are unknown). leaves mostly ternate, linear-subulate,(4–)6–10(–12)
Grazing by goats prevents successful regeneration × 0.5–0.7 mm, pungent, adaxially canaliculate, epis-
and it is hardly possible to speak of viable popula- tomatic, with two narrow bands of stomata. Scale
tions in most places. leaves increasingly dominant with age of plant in 395
IUCN: EN (A2c, D) most varieties, always decussate, imbricate, closely
appressed, on ultimate branchlets 1.5–3 × 1 mm,
ovate-rhombic to rhombic-lanceolate, abaxially
Uses
with a conspicuous central, elliptic gland, green
When Canary Island juniper was still abundant its or glaucous green; margins entire; apex obtuse to
wood was highly sought after due to its durability; acute; stomata few at base abaxially, in two concave
it was used for fence posts and for furniture mak- fields separated by a narrow midrib adaxially. Pollen
ing. This species is rare in cultivation and ought to cones terminal on ultimate branchlets with scale-
be tried more (perhaps not in countries with rela- like or acicular leaves, ovoid-oblong, 4–6 × 2–3 mm;
tively few hot, sunny days in summer). It grows on microsporophylls 12–18, decussate, peltate-cordate,
Tenerife at altitudes where frost and snow are not convex, upper margin entire, abaxially bearing 3–4
uncommon, but solar radiation is intense. Ex situ ovoid-oblong pollen sacs. Seed cones terminal on
conservation is urgently needed to build up stock erect, short shoots 3–10 mm long with small scale
for future replanting and horticulture could have an leaves, maturing in second season to (irregularly)
important role to play in this effort. globose or broad pyriform, 4–10 mm, usually dry
pulpy, resinous, light brown, red-brown or glaucous
cones. Bract-scale complexes (4–)6(–8), decussate,
Juniperus chinensis L., Mant. Pl. 1: 127. 1767. occasionally ternate, usually the lower pair infertile,
entirely fused, sutures of upper scales partly visible
as a curved ridge terminating in a minute bract tip.
Etymology
Seeds (1–)2–4(–5) per cone, ovoid-flattened, 3–7
The species epithet means “from China”. × 2–6 mm, with small resin pits and longitudinal,
shallow grooves, light yellowish brown.
Vernacular names
Taxonomic notes
Chinese juniper; yuan bai (Chinese); Ibuki, Byaku­
shin, Ibuki-byakushin, Kamakura-byakushin (Japa- Juniperus chinensis is one of the most variable spe-
nese); Kong Nam Tsong (Korean) cies in the genus. Its polymorphy has been the cause
of an almost inextricable synonymy, especially so
because horticulturists have described taxa at vari-
Description
ous ranks from variations observed in cultivation
Decumbent, sometimes ascending or erect shrubs, which should have been treated as cultivars.
or a tall trees to 20–25 m with a monopodial, erect
trunk to 1 m diam., dioecious or rarely monoecious.
Distribution
Bark on tree trunks fissured, exfoliating in long, fi-
brous, thin strips, becoming darker grey-brown. Myanmar [Burma], China, Japan, Korea, Russian
Branches spreading horizontally or ascending, in Far East, Taiwan.
shrubs spreading, ascending or erect, spreading TDWG codes: 31 KHA KUR PRM SAK 36 CHC-CQ
or subpendulous in trees, forming dense matting CHC-GZ CHC-HU CHC-SC CHC-YN CHI-NM CHI-NX
crowns in decumbent shrubs to initially pyramidal, CHM-HJ CHM-JL CHM-LN CHN-BJ CHN-GS CHN-HB
 CHN-SA CHN-SD CHN-SX CHN-TJ CHS-AH CHS-FJ This species is also of major importance in bonsai
CHS-GD CHS-GX CHS-HE CHS-HK CHS-HN CHS- and penjing culture.
JS CHS-JX CHS-ZJ CHT 38 JAP-HK JAP-HN JAP-KY
JAP-SH KOR-NK KOR-SK TAI 41 MYA 3 varieties are recognized:

Juniperus chinensis L. var. chinensis. Type: Sweden:

Ecology
[cultivated in Uppsala Bot. Garden, Sweden], leg.
In a few localities this widespread species forms ign. LINN 1198.3 (lectotype LINN).
groves of tall trees (e.g. in S Gansu), or it is mixed
with pines and deciduous angiosperms at canopy Sabina vulgaris Antoine var. erectopatens
396 level. It is much more common, under conditions W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13
largely determined by man’s agricultural practices, (4): 86. 1975; Juniperus sabina L. var. erectopatens
in secondary vegetation, on open, rocky slopes. The (W. C. Cheng & L. K. Fu) Y. F. Yu & L. K. Fu, Novon
altitudinal range is (100–)1400–2400(–2700) m a.s.l. 7 (4): 444. 1998; Juniperus erectopatens (W. C. Cheng
Widespread planting and subsequent establishment & L. K. Fu) R. P. Adams, Biochem. Syst. Ecol. 27 (7):
in areas where it was not originally native have made 723. 1999.
it difficult to establish it original habitat and types
of vegetation. While now predominantly montane,
Description
it may have been part of lowland forests in the past
(Wang, 1961). High montane varieties J. chinensis Erect shrubs to trees. Leaves of two kinds, needle-
var. sargentii and var. tsukusiensis occur on rocky like and scale-like, present on a single plant; needle-
outcrops and amongst boulders and have attained a like leaves mostly in whorls of 3, rarely decussate,
decumbent habit. 6–12 mm long; scale leaves decussate. Pollen and
seed cones only on ultimate branchlets with scale
leaves.
Uses
Juniperus chinensis is (with Platycladus orientalis)
Distribution
one of the two most commonly planted cupressa-
ceous trees in traditional Chinese gardens, such as Myanmar [Burma]; China; Japan; Korea; Taiwan.
around temples and in the extensive grounds of the TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
Forbidden City in Beijing. These grounds are now SC CHC-YN CHI-NM CHN-BJ CHN-GS CHN-HB
virtual reserves for large specimen trees, which have CHN-SA CHN-SD CHN-SX CHN-TJ CHS-AH CHS-FJ
all but disappeared from the countryside. Its wood CHS-GD CHS-GX CHS-HE CHS-HK CHS-HN CHS-JS
is highly valued for furniture making and joinery CHS-JX CHS-ZJ 38 JAP-HK JAP-HN JAP-KY JAP-SH
and is hard and durable. In horticulture, J. chinen- KOR-NK KOR-SK TAI 41 MYA
sis has been the source of many cultivars and (as
J. ­sphaerica Lindl.) one parent of a purported hy-
Conservation
brid (of garden origin) with J. sabina, for which Van
Melle (1946) proposed the name J. ×media, a later IUCN: LC
homonym of J. media V. D. Dmitriev (1938) = J. semi­
globosa. A proposal to conserve van Melle’s name Juniperus chinensis L. var. sargentii A. Henry,
failed and the cultivars thought to derive from this in Elwes & Henry, Trees Gr. Brit. Ireland 6: 1432.
hybrid origin are now to be listed under J. ×pfitzeri- 1912. Juniperus sargentii (A. Henry) Takeda ex
ana based on a selection made by the Späth Nursery Nakai, Bot. Mag. (Tokyo) 44: 511. 1930. Type: Japan:
in Berlin, Germany in the 1890’s (Pfitzer Junipers). Hokkaido, Kitami Prov., Rebun-jima, M. Furuse
A study using RAPDs (Le Duc, Adams & Zhong, 9385 (neotype K). Fig. 134
1999) supports the notion of a hybrid origin with
parents J. chinensis and J. sabina of the Pfitzer culti-
Vernacular names
vars, for which the correct botanical name is Junipe-
rus ×pfitzeriana (Späth) Schmidt [‘Pfitzer Group’]. yan bai (Chinese); Miyama-byakushin (Japanese)
 thick, quadrangular in cross-section, lax but not
Description
drooping. Leaves on most shrubs of two types, nee-
Decumbent shrubs, forming broad, dense mats. dle-like and scale-like, or only scale leaves; acicular
Branches ascending, whip shoots assurgent, later leaves in whorls of 3, 4–8 mm long; scale leaves 1–1.3
prostrate and rooting. Leaves all needle-like only on × 1 mm, triangular rhombic or gibbous, obtuse or
seedlings and young plants, gradually replaced by slightly apiculate. Gland central, small, ovoid, con-
scale leaves on mature plants (but sometimes still spicuous but inactive. Seed cones small 4–5 mm
present on coning branches), all decussate, the aci­ diam., red-brown to purplish brown, with some
cular leaves 3–7 mm long. glaucous bloom.

397
Distribution Distribution
China: Heilongjiang; Japan, Russian Far East. China?, Taiwan (Chingshuei Cliffs), S Japan (Yaku­
TDWG codes: 31 KHA KUR PRM SAK 36 CHM-HJ 38 shima).
JAP-HK JAP-HN JAP-KY JAP-SH TDWG codes: 38 JAP-KY TAI

Ecology Ecology
This variety is a constituent of high montane to This variety has been found in a few localities in
subalpine scrubland and meadows, although in the high mountains as a procumbent shrub but little
far north of its range (Sakhalin Island) it occurs at else is know about its ecology. In Taiwan it is found
much lower altitude. on steep cliffs high above the Pacific Ocean, where
its stems spread over bare rock.
Conservation
Conservation
IUCN: LC
Known from Yakushima and three localities in the
central mountains of Taiwan, this taxon is undoubt-
Uses
edly rare, but we do not know its entire distribution
This dwarfed form of the species has obvious horti- (it may have been overlooked as most collections in
cultural merits and is cultivated in many countries Taiwan are relatively recent) to make a reliable eval-
and several cultivars exist of it. It was introduced uation of its conservation status.
to the Arnold Arboretum around 1892 by the first IUCN: DD
­Director, Charles S. Sargent.

Juniperus chinensis L. var. tsukusiensis (Masam.) Juniperus coahuilensis (Martínez) Gaussen ex

Masam., J. Soc. Trop. Agric. Formosa 2: 152. 1930 R. P. Adams, Phytologia 74: 413. 1993. Juniperus
[& in 3: 20. 1931]. Juniperus tsukusiensis Masam., erythrocarpa Cory var. coahuilensis Martínez,
Bot. Mag. (Tokyo) 44: 50. 1930. Type: Japan: Anales Inst. Biol. Univ. Nac. México 17 (1): 114.
Kyushu, Yakushima, G. Masamune s.n. (holotype 1946. Type: Mexico: Coahuila, Sierra de los Hechi­
not designated, syntype TI). ceros, Cañon de La Madera, I. M. Johnston &
C. H. Mueller 1290 (holotype MEXU).
Vernacular names
Etymology
qing shui yuan bai (Chinese)
The species epithet refers to the Mexican State of
Coahuila.
Description
Decumbent or low shrubs to 80–100 cm, often pros-
Vernacular names
trate or creeping over rocks. Foliage very dense, ul-
timate branchlets 8–25 mm long, slender, ca. 1 mm No common names are recorded for this species.
 Description Ecology
Large shrub to 5 m, or small tree to 10 m, dioecious, This species occurs in the high desert (1200–2000
rarely monoecious; multistemmed or with a short m a.s.l.) Bouteloua grasslands and adjacent rocky ar-
trunk and a maximal diam. of 50 cm. Bark of thick eas. It is unusual, for Juniperus, in that it occurs in
branches and trunk fibrous, exfoliating in strips, these seemingly undisturbed grasslands. In Mexico,
weathering ash-grey. Branches numerous, long, the populations may occur in canyons or on alluvial
ascending from base or spreading, often curved, fans and, of course, it is found in rocky areas adja-
forming a broad, rounded or more irregular and cent to grasslands, often in association with Opuntia
open crown. Foliage branches numerous, irregu- spp. or Yucca spp. It is unusual in that J. coahuilensis
398 larly disposed but not pendulous, ultimate branch- sprouts from the stump when cut or burned. This
lets spreading 30–70 degrees, slender and stiff, up feature has probably allowed it to remain in the
to 15 mm long, 1–1.5 mm thick, (weakly) quadran- grasslands in spite of periodic grass fires that kill all
gular in cross section, covered with scale leaves, other juniper species [R. P. Adams, pers. comm.]. It
persistent. Leaves decussate on all ultimate branch- is considered to be ‘weedy’ in some areas by sheep-
lets, decussate or in alternate whorls of 3 on older men and cattlemen.
branchlets, imbricate, scale-like, 1.2–1.7(–1.8) × 0.7–
1.2 mm on lateral branchlets, decurrent, rhombic or
Conservation
ovoid-rhombic; apex appressed or rarely free, obtuse
to acute; margins minutely denticulate; stomata on IUCN: LC
abaxial side limited to two small spots near leaf base,
on adaxial surface in two tapering bands; glands
central on small scale leaves, more or less conspicu- Juniperus comitana Martínez, Anales Inst. Biol.
ous, large, rounded to oblong, flat or slightly raised, Univ. Nac. México 15 (1): 12. 1944. Type: Mexico:
exudate often present; leaf colour yellowish green or Chiapas, Comitán de Domínguez, 12 km S of
light green. Pollen cones numerous, terminal, soli- Comitán, M. Martínez 6700 (holotype MEXU).
tary, ellipsoid or ovoid-oblong, 3–4 × 2–2.5 mm; mi-
crosporophylls 8–10, decussate, peltate, acute, with
Etymology
minutely denticulate upper margins and bearing 3–4
abaxial pollen sacs. Seed cones terminal, maturing The species epithet refers to the town of Comitán in
in one season attaining variable colours from pink- Chiapas, Mexico.
ish(–red) or orange(–red) to purplish blue, usually
glaucous, (sub-)globose or sometimes broad ovoid,
Vernacular names
5–7 mm diam. (if ovoid to 9 mm long), internally
soft pulpy, succulent, yellowish to orange-brown. No common names have been recorded for this spe-
Bract-scale complexes 4(–6), entirely fused, decus- cies.
sate; bract apices minutely exserted or hidden; sur-
face smooth, rugose when dry. Seeds 1(–2) per cone,
Description
ovoid-globose, 3–5(–6) × 3–4(–5) mm, with rounded
base and acutish apex, shallowly grooved and ridged Trees to 8–10 m, presumably dioecious; trunk
on sides, not or slightly resinous, chestnut brown, monopodial, erect or forked and twisted, usually
with a large, lighter lobed hilum ­proximally. branching well above base, up to 60–80 cm d.b.h.
Bark on trunk 5–10 mm thick, fibrous, lacerated, ex-
foliating in long strips exposing purplish inner bark;
Distribution
outer bark red-brown to grey-brown. Branches long,
N and NE Mexico: Chihuahua, Coahuila; USA: SW spreading or ascending, foliage forming in mature
Texas. trees a broad, rounded or irregular crown. Foliage
TDWG codes: 77 TEX 79 MXE-CO MXE-CU branches numerous, forming lax, dense foliage tufts,
irregularly disposed, ultimate branchlets of ­irregular
length from 4–20(–25) mm, very slender, 0.7–1 mm
diam., more or less quadrangular in cross-section to Sierra de los Cuchumatánes where both J. comitana
more terete on older branchlets which become rough and J. standleyi still occur.
with free leaf apices, persistent. Leaves on ultimate IUCN: VU (B1+2c)
and lower lateral branchlets decussate, but occasion-
ally on slower growing sections of older branchlets
Uses
in alternating whorls of 3, closely appressed, on ulti-
mate branchlets ovoid-rhombic or rhombic-triangu- The wood of this juniper is used locally by the
lar, 1–1.4 × 0.7–1 mm; margins ­hyaline and minutely ­Amerindian population for firewood and also for
erose to serrulate; apex acuminate or sometimes fence posts.
obtuse; stomata few abaxially near base along mar-
gins, abundant adaxially from base to apex; glands 399
small, usually inactive; leaf colour light green. Pol- Juniperus communis L., Sp. Pl. 2: 1040. 1753.
len cones terminal, solitary, oblong, 4–6 mm long;
microsporophylls 10–12, decussate, peltate, with
Etymology
minutely hyaline-denticulate upper margins, bear-
ing (3–)4 abaxial pollen sacs. Seed cones numerous, The species epithet means “common”; Linnaeus
terminal, solitary on very short lateral branchlets, knew it in abundance from Sweden.
maturing in one year to purplish brown cones with a
glaucous bloom; mature cones subglobose, 5–7 mm
Vernacular names
diam. Bract-scale complexes 4, decussate, lower pair
smaller than upper pair meeting at cone apex, com- Common juniper; Genévrier commun (French);
pletely fused, with smooth external surface, lustrous Gemeine Wacholder (German); Mozhzhevel’nik
when ripe; 2 decussate pairs of bract tips visible but obyknovennyy (Russian)
these less than 1 mm long, in ripe cones sometimes
submerged; internal tissue soft pulpy, resinous.
Description
Seeds usually 1 per cone, ovoid, 4–5(–6) × 3.5–4 mm.
Decumbent, spreading or large shrubs to small
trees, to 12 m tall, dioecious, rarely monoecious;
Distribution
multistemmed or occasionally monopodial (trunk
Mexico: Chiapas (Comitán); N Guatemala. to 2–2.5 m tall, diam. to 50 cm), low branching.
TDWG codes: 79 MXT-CI 80 GUA Bark on large trunks becoming fissured, with thin
strips, grey-brown. Branches numerous, decum-
bent, spreading, ascending or erect. Foliage usually
Ecology
dense and stiff, but sometimes more open and lax
Juniperus comitana is found on dry, rocky slopes to subpendulous, with numerous ‘jointed’ ultimate
or in canyons with shrub or open woodland cover, branchlets, persistent, forming a very variable crown
growing with e.g. Acacia and Ficus in forest pasture (habit) largely influenced by site conditions. Leaves
on dolomite and other rock types with thin soil. The in alternating whorls of 3, rarely 4, non-decurrent,
altitudinal range is from 1300 m to 1800 m a.s.l. spreading 30–90 degrees at nodes 2–15 mm apart,
rigid, slightly thickened at base, jointed to leaf part
adnate with shoot, (narrowly) linear to broadly lan-
Conservation
ceolate (‘boat-shaped’), 4–25 × 0.8–2.4 mm, straight
This species is threatened by deforestation and over- or curved-falcate; abaxial face keeled or with a faint
exploitation of forest resources as an indirect result of midrib, green, sometimes glaucous green; adax-
a rapidly growing human population with an almost ial margins narrow or broad relative to the single
exclusively rural economy. Especially the junipers stomatal band, entire; apex obtuse to acuminate or
that occur in pine-oak forests, although not specifi- pungent; stomata in a conspicuous band bordered
cally targeted, have decreased with that forest type by green margins. Pollen cones axillary to leaves,
both in Chiapas and in Guatemala. The situation in solitary, 1–3 per leaf whorl, subglobose to ellipsoid-
Guatemala has been described by Islebe (1993) with oblong, 3–5 mm long; microsporophylls 9–12, in
some emphasis on the situation in the Guatemalan alternating whorls of 3, triangular-peltate, with
 erose-denticulate margins and acute apex, bearing KOR-SK 40 NEP PAK WHM-HP WHM-JK WHM-UT
3–6 abaxial pollen sacs near lower margin. Seed 70 ASK GNL NUN NWT YUK 71 ABT BRC MAN SAS 72
cones axillary on very short (1 mm) dwarf shoots LAB NBR NFL-NE NFL-SP NSC ONT PEI QUE 73 COL
with whorls of minute scale leaves, maturing in IDA MNT ORE WAS WYO 74 ILL IOW KAN MIN MSO
18 months; mature cones globose or broadly ovoid, NDA NEB OKL SDA WIS 75 CNT INI MAI MAS MIC
4–13 mm diam., with 1(–2) whorls of 3 completely NWH NWJ NWY OHI PEN RHO VER WVA 76 ARI CAL
fused bract-scale complexes, only the upper whorl NEV UTA 77 NWM 78 GEO KTY NCA SCA TEN VRG
fully developed, sutures not visible, bract tip hidden
or very small; tissue soft pulpy, juicy in most cones;
Ecology
mature cones from purplish red to black-brown, of-
400 ten with dark glaucous blue bloom. Seeds 1–3 per This most widespread of all conifer species occurs in
cone, (tri)angular, oblong, (3–)4–5 × 2–3 mm, light a very wide range of habitats, from lowland bogs and
brown with shallow pits and acute apex. coastal sands to high alpine meadows, moraines, and
scree slopes. A limited number of geographical vari-
eties is here recognized, some are ecologically more
Taxonomic notes
restricted than the species as a whole, but there is
Juniperus communis is a polymorphic species. In al- considerable overlap. More detailed information is
most all cases infraspecific taxa that have been de- given under these varieties.
scribed are either largely sympatric or merge into
each other morphologically where they meet. Habit
Uses
(growth form) is largely dependent on habitat fac-
tors, foremost of which is climate (temperature, The Common juniper is widely used as an ornamen-
precipitation, duration of snow cover) and if not sup- tal in parks and gardens, especially the decumbent
ported by other characters it is unlikely to indicate varieties, and numerous cultivars have been derived
distinct taxa. The recognition of a two-ranked hier- from it. The cones (‘berries’) are used in cooking and
archy of infraspecific taxa is especially inappropriate in the preparation of alcoholic distilled beverages.
in this species where within-population variation Juniperus communis var. communis has been used for
can be of the same order (measuring morphology) a considerable time in European horticulture, where
as that between recognized infraspecific taxa. especially the fastigiate habit of some NW European
plants is popular, and a number of cultivars have
been obtained by selection. Juniperus communis var.
Distribution
saxatilis, being a prostrate shrub, is a useful plant in
Temperate Eurasia, North America N of Mexico. horticulture, where it is usually known as J. commu-
Juniperus communis is the most widely distributed nis var. montana or var. nana, or under a number
conifer species in the world, with a circumpolar dis- of cultivar names (= vegetatively propagated clones)
tribution extending from ca. 70° N in Alaska, Scan- selected for dwarfed growth, branching habit, foli-
dinavia, and Siberia to ca. 28° N in the Himalaya. age colour, etc.
TDWG codes: 10 DEN FIN FOR GRB ICE IRE NOR
SWE 11 AUT-AU AUT-LI BGM-BE BGM-LU CZE-CZ 5 varieties are recognized:
CZE-SK GER HUN NET POL SWI 12 BAL COR FRA-
CI FRA-FR POR SAR SPA-AN SPA-GI SPA-SP 13 ALB Juniperus communis L. var. communis. Type:
BUL GRC ITA-IT KRI ROM SIC-SI TUE YUG-BH YUG- Europe, Alps?, leg. ign. HSC s.n. (lectotype BM).
CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 Fig. 135
BLR BLT-ES BLT-KA BLT-LA BLT-LI KRY RUC RUE
RUN RUS RUW UKR-MO UKR-UK 30 ALT BRY CTA
Description
IRK KRA TVA WSB YAK 31 AMU KAM KHA KUR MAG
PRM SAK 32 KAZ KGZ TKM TZK UZB 33 NCS TCS 34 Shrubs or small trees to 10–12 m with variable habit,
AFG CYP EAI IRN IRQ LBS-LB LBS-SY TUR 36 CHI- often conical-pyramidal but also spreading, round-
NM CHM CHX 37 MON 38 JAP-HK JAP-HN KOR-NK ed, dense or open, or a tree with short trunk 2–2.5
m tall and a pyramidal, rarely flat-topped crown. Juniperus communis L. var. depressa Pursh, Fl.
Leaves spreading at nodes (2–)5–10(–15) mm apart Amer. Sept. 2: 646. 1814. Juniperus depressa (Pursh)
and at wide or right angle to shoot, (4–)7–20(–25) × Raf., Med. Fl. 2: 13. 1830; Juniperus communis
0.7–1.5 mm, linear, straight, tapering above the mid- L. subsp. depressa (Pursh) Franco, Bol. Soc. Brot.,
dle to a pungent tip. Leaf length variable between ser. 2, 36: 117. 1962. Type not designated.
individual plants as well as on single plants, usually
more than twice as long as mature cones. Seed cones
Vernacular names
globose or broadly ovoid, (4–)5–7(–8) mm, black-
brown, with blue bloom, or purplish black. Seeds 3 Dwarf juniper, Prostrate juniper
per cone.
401
Description
Distribution
Decumbent or spreading shrubs to 1.5 m, rarely
Europe, North Africa, Caucasus, Siberia to Russian a large shrub or small tree to 10 m, not forming a
Far East, Western Asia, Central Asia. conical or pyramidal crown; branchlets ascending at
TDWG codes: 10 DEN FIN FOR GRB ICE IRE NOR tips. Leaves at nodes 3–6(–10) mm apart, spreading
SWE 11 AUT-AU AUT-LI BGM-BE BGM-LU CZE-CZ at angles of 30–70 degrees, often ascending or more
CZE-SK GER HUN NET POL SWI 12 BAL COR FRA-CI or less erect, at internodes of c. 5 mm, curved at base
FRA-FR POR SAR SPA-AN SPA-SP 13 ALB BUL GRC (sometimes straight), linear, (5–)10–20 × 1–1.7 mm;
ITA-IT ITA-SM KRI ROM SIC-SI TUE YUG-BH YUG- stomatal band from as wide as each green margin to
CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 nearly twice as wide. Seed cones 6–9 mm, globose.
BLR BLT-ES BLT-KA BLT-LA BLT-LI KRY RUC RUE Seeds 3 per cone.
RUN RUS RUW UKR-MO UKR-UK 30 ALT BRY CTA
IRK KRA TVA WSB YAK 33 NCS TCS 34 AFG CYP EAI
Distribution
IRN IRQ LBS-LB LBS-SY TUR
North America, from Alaska to Newfoundland, S to
Arizona and South Carolina.
Ecology
TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
This, the ‘typical’ variety, is largely a pioneer wood- MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT PEI
land species, occupying natural rock outcrops and QUE 73 COL IDA MNT ORE WAS WYO 74 ILL IOW
other places with skeletal soil and abundant sunlight KAN MIN MSO NDA NEB OKL SDA WIS 75 CNT INI
in woodland and light forest, both broad-leaf and MAI MAS MIC NWH NWJ NWY OHI PEN RHO VER
coniferous forest (especially Pinus sylvestris-­Betula WVA 76 ARI CAL NEV UTA 77 NWM 78 GEO KTY
spp.-Quercus spp.), in which it can obtain local NCA SCA TEN VRG
dominance after disturbances (non-fire). It is also
prevalent in the ecotone between open woodland
Ecology
and grassland on poor sandy soils and on stabilised
inland sand dunes. It occurs often with Calluna vul- In a wide range of habitats from sea shores to (sub)
garis, Erica spp., Vaccinium spp., Arbutus sp., Cytisus alpine meadows and rocky ridges. Mostly associated
scoparius, Ulex sp., Salix spp., and the above men- with coniferous woodland and forest, but usually
tioned tree genera, in Russia also in grass steppes. occupying localities or topography less suitable for
The altitude ranges from 5 m to 2400 m a.s.l. It tree growth, or in open woodland on thin or poor
seems very indifferent to soil type and occurs in dry sandy soil or dolomite. Associated e.g. with Pinus
sand, chalk downs, and loose (dolomitic) scree, as banksiana, P. ponderosa, P. flexilis, Abies balsamea,
well as in acidic peat, with low or fluctuating ground A. lasiocarpa, Picea engelmannii, P. glauca, and Popu-
water levels. lus tremuloides in lowland to subalpine forests, with
shrubs and herbs in coastal or inland sand dunes, in
(sub)alpine meadows, or in peat bogs. The altitudinal
Conservation
range is from 5 m to 3800 m a.s.l. It is indifferent to
IUCN: LC soil types, with pH from very acid to neutral or mild-
ly alkaline and with low or fluctuating water tables.
 Conservation Description
IUCN: LC Decumbent or spreading shrubs 0.5–1.5 m tall with
spreading branches. Leaves spreading at nodes 3–5
Juniperus communis L. var. megistocarpa Fernald mm apart, ascending, acicular, 7–10 × 0.8–1 mm,
& H. St. John Type: Canada: Quebec, Madeleine usually distinctly curved (falcate), triquetrous; adax-
Islands, Alright Island, Narrows, M. L. Fernald & ial face sulcate; stomatal band narrow, nearly hid-
B. H. Long 6729 (holotype GH). den from view; apex pungent. Seed cones globose
or ovoid-globose, 4–6(–7) mm, brown-pruinose to
blue-black. Seeds 1–3 per cone.
Description
402
Decumbent shrubs, often creeping over rock or
Distribution
sand. Leaves at nodes 2–5 mm apart, ascending,
short and wide, 4–12 × 1.6–2.4 mm, more or less Japan: Hokkaido, N Honshu.
boat-shaped or broad-linear, acuminate-pungent, TDWG codes: 38 JAP-HK
with broad stomatal band slightly wider than one of
the broad green margins. Seed cones often as large
Ecology
or larger than leaves, 9–13 mm, soft pulpy, blackish
blue or glaucous blue. Seeds 3 per cone. Insufficiently known. The known altitudinal range is
230–650 m a.s.l.
Distribution
Conservation
E. Canada: Newfoundland, Nova Scotia (Sable Is-
land), Quebec (Île de la Madeleine). IUCN: LC
TDWG codes: 72 NFL-NE NFL-SP NSC QUE
Juniperus communis L. var. saxatilis Pall., Fl.
Rossica 1 (2): 12. 1789. Type: Illustration in Pallas,
Ecology
Fl. Rossica 1 (2): 12, t. 54. 1789 (lectotype). Fig. 136
In Empetrum heath and open vegetation on sand
dunes and on serpentine and magnesian limestone Juniperus sibirica Burgsd., Anleit. Sich. Erzieh. Hol-
barrens, commonly in the ecotone between open zart. 2: 124. 1787.
vegetation and coniferous forest with Abies balsa- Juniperus communis L. var. montana Aiton, Hort.
mea or Picea rubens. Kew. 3: 414. 1789; Juniperus montana (Aiton) Lindl.
& Gordon, J. Hort. Soc. London 5: 200. 1850.
Juniperus pygmaea K. Koch, Linnaea 22: 302. 1849;
Conservation
Juniperus communis L. subsp. pygmaea (K. Koch)
IUCN: LC Imkhan., Novosti Sist. Vyssh. Rast. 27: 10. 1990.
Juniperus communis L. var. charlottensis R. P. Ad-
Juniperus communis L. var. nipponica (Maxim.) ams, Phytologia 90 (2): 187. 2008.
E. H. Wilson, [Conifers & Taxads Japan] Publ. Ar-
nold Arbor. 8: 81. 1916. Juniperus nipponica Maxim.,
Vernacular names
Bull. Acad. Imp. Sci. Saint-Pétersbourg 12: 230. 1868;
Juniperus rigida Siebold & Zucc. subsp. nipponica Mountain juniper; Zwergwacholder (German);
(Maxim.) Franco, Bol. Soc. Brot., ser. 2, 36: 119. 1962. Ginepro nano (Italian); Genévrier nain (French);
Type: Japan: Honshu, Iwate Pref., [“Prov. Nambu, xian bei ci bai (Chinese)
in alpibus altioribus”], Tschonoski (Chônosuka Sug-
awa) [ex herb. C. J. Maximowicz s.n.] (holotype LE).
Description
Decumbent or spreading shrubs with spreading
Vernacular names
branches. Leaves at nodes 2–4 mm apart, spread-
Miama-nezu (Japanese) ing or ascending, often almost imbricate, straight or
curved, 4–10 × 1–2 mm, flattened-concave, keeled outcrops in sunny, relatively dry, early snow-free
abaxially; apex acuminate-pungent; stomatal band expositions at (sub)alpine altitudes; also abundant
wider than green margins. Seed cones ovoid-glo- in Pinus mugo thickets and as undergrowth in mon-
bose 4–7(–10) mm, brown-pruinose, blue or blue- tane, open Larix and Pinus forests, in Central Asia
black. Seeds 1–3 per cone. also with Picea schrenkiana and Abies sibirica, or
with Betula sp., elsewhere (e.g. Caucasus, Himalaya,
Rocky Mountains) with other species of Abies, Picea
Taxonomic notes
or Pinus. Also on lake shores and in peat bogs in
The correct name of this taxon under Juniperus com- mountains, associated with typical holarctic plants
munis has not been agreed upon in the horticultural e.g. Vaccinium, Arctostaphylos, Cotoneaster, Rosa,
and taxonomic literature, where the epithets ‘alpina’, Artemisia (the latter especially in Asian mountains 403
‘montana’, ‘nana’ and ‘saxatilis’ are all to be found in and steppes), and grasses. Mostly on acidic soils but
use for what is evidently the same plant. All four and in dry places also on limestone.
J. sibirica Burgsd. have been used at specific rank as
well, and some in addition at subspecific rank, while
Conservation
the nomenclature has been further confused by the
adoption of various combinations of these epithets. IUCN: LC
Adams (op. cit.) recently considered that this taxon
does not occur in Japan, Sakhalin and Kamchatka
(all treated as var. nipponica), and that the shrubs Juniperus convallium Rehd. & E. H. Wilson, in
of this species on the coast of NW North America Sargent, Pl. Wilson. 2: 62. 1914.
should be recognized as yet another variety, J. com-
munis var. charlottensis. Morphologically, at least,
Etymology
these plants are all very similar.
The species epithet probably derives from Latin con-
vallis = ‘deep, enclosed valley’, and if so, is referring
Distribution
to its habitat.
Europe, Caucasus, Siberia, Central Asia, W Asia,
Jammu-Kashmir, Himachal Pradesh, Uttar Pradesh,
Vernacular names
Nepal, Pakistan, NE & NW China, Japan, Korea,
Russian Far East, W North America, Greenland. mi zhi yuan bai (Chinese)
TDWG codes: 10 NOR SWE 11 CZE-CZ CZE-SK GER
POL SWI 12 COR FRA-FR SPA-AN SPA-SP 13 ALB BUL
Description
GRC ITA-IT KRI ROM SIC-SI TUE YUG-BH YUG-
CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 Small to medium size trees (rarely shrubs) to 20
UKR-MO UKR-UK 30 ALT BRY CTA IRK KRA TVA m, dioecious or monoecious; trunk multistemmed
WSB YAK 31 AMU KAM KHA KUR MAG PRM SAK 32 or monopodial, up to 50 cm d.b.h. Bark on larger
KAZ KGZ TKM TZK UZB 33 NCS TCS 34 AFG CYP EAI stems exfoliating in longitudinal strips, grey-brown.
IRN IRQ LBS-LB TUR 36 CHI-NM CHM CHX 37 MON Branches spreading or ascending, foliage branches
38 JAP-HK KOR-NK KOR-SK 40 NEP PAK WHM-HP very dense, short, stiff and speading, or longer and
WHM-JK WHM-UT 70 ASK GNL 71 BRC 73 ORE WAS more or less lax, forming a dense, rounded crown.
76 CAL Foliage branchlets ultimately slender, subterete or
weakly quadrangular, 0.8–1.2 mm wide, covered
with closely appressed leaves, persistent. Leaves on
Ecology
mature plants scale-like, decussate (sometimes ter-
Widely distributed in montane to (sub)alpine veg- nate), imbricate, decurrent, 1.5–2 × 0.8–1 mm, rhom-
etation types of high mountains in Eurasia and bic, keeled near apex or not, obtuse; epistomatic,
western North America; also in the Asian high stomata in a proximally divided band; leaves abaxi-
steppes. The altitudinal range is great: from 5 m to ally glandular (in one variety often inconspicuous);
4050 m a.s.l. In the Alps forming dominant veg- gland concave or convex, yellowish or darker than
etation cover over usually siliceous scree and rock the light green or greyish green leaf. Pollen cones
 numerous, solitary, terminal on short branchlets, 2 varieties are recognized:
globose to subglobose, 2–3 mm; microsporophylls
decussate, 6–8, peltate-triangular, with obtuse apex, Juniperus convallium Rehd. & E. H. Wilson var.
bearing 2–3 abaxial pollen sacs near the lower mar- convallium. Type: China: Sichuan, NW Sichuan,
gin. Seed cones terminal on short, curved or erect (loc. not indicated), Wilson 3010 (holotype A).
branchlets, maturing in the second season to sub-
globose or ovoid-conical, 5–8(–10) × 5–6 mm, light
Description
reddish brown to purplish black or glaucous cones.
Bract-scale complexes 4–6, decussate, entirely fused, Leaves conspicuously glandular, not keeled near
sometimes sutures faintly visible near apex of cone; apex. Seed cones subglobose to ovoid-conical, 6–8
404 bract tip usually hidden or barely visible; scale tis- (–10) × 5–6 mm. Seeds 4–5 mm diam., subglobose
sue dry pulpy or more succulent. Seeds 1 per cone, to globose-conical or ovoid-conical.
subglobose (onion-shaped) or globose-conical to
ovoid-conical, sometimes slightly flattened, 3–5 mm
Distribution
diam., shallowly grooved or bifacially ridged, with
small resin pits near base. China: S Gansu, SE Qinghai, W Sichuan, E Xizang
[Tibet], NW Yunnan.
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHQ
Distribution
CHT
China: S Gansu, SE Qinghai, W Sichuan, E Xizang
[Tibet], NW Yunnan.
Conservation
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHQ
CHT IUCN: LC

Juniperus convallium Rehd. & E. H. Wilson var.

Ecology
microsperma (W. C. Cheng & L. K. Fu) Silba, Phy-
In high montane to subalpine coniferous forest, juni- tologia Mem. 7: 33. 1984. Sabina convallium (Rehd.
per woodland, occasionally in the ecotone to alpine & E. H. Wilson) W. C. Cheng & L. K. Fu var. micro-
steppe, at high altitudes often on S-SW facing slopes sperma W. C. Cheng & L. K. Fu, Acta Phytotax. Sin.
or on sun-warmed rock outcrops, both of limestone 13 (4): 86. 1975; Sabina microsperma (W. C. Cheng
and granitic rock. The altitudinal range is 2200– & L. K. Fu) W. C. Cheng & L. K. Fu, Fl. Xizangica 1:
4430 m a.s.l. Associated with Abies, Picea (these 390. 1983; Juniperus microsperma (W. C. Cheng &
genera predominantly on N-NE facing, moister and L. K. Fu) R. P. Adams, Biochem. Syst. Ecol. 28: 540.
cooler slopes), Juniperus saltuaria, J. tibetica (which 2000. Type: China: Xizang (Tibet), Mekong River,
usually reaches higher altitudes), Quercus aquifolio- Qamdo, [“Chang Du”], leg. ign. [“(First) Exped. to
ides, Seriphidium (Artemisia), Caragana, and Rosa; Xizang (Tibet)”] 10019 (holotype PE).
woodlands are often degraded to scrub-pasture. In
the dryer valleys beyond the direct monsoon influ-
Vernacular names
ence the lower limit of annual rainfall (mostly sum-
mer) is ca. 300 mm at 3600–3800 m. Low winter xiao zi yuan bai (Chinese)
temperatures, to which this species is more sensitive
than J. tibetica, cause it to remain below the latter in
Description
many areas, but there are mixed stands especially on
S-SW exposed slopes where J. convallium can occur Leaves often inconspicuously glandular, keeled near
at higher altitudes. apex. Seed cones ovoid, 5–7 × 5 mm. Seeds 4 × 3
mm, slightly flattened-ovoid.
Uses
Distribution
Not known, probably for firewood in areas with few
trees. China: W Sichuan, SE Xizang [Tibet].
TDWG codes: 36 CHC-SC CHT
 apex, bearing 3–5 abaxial angular pollen sacs. Seed
Conservation
cones terminal on very short, erect lateral branch-
The distribution of this form seems to be more lim- lets, maturing in the second season to (sub)globose
ited than that of the species as a whole, but its extent or broadly ovoid, 7–15 mm diam., light or red-
remains insufficiently known, in part due to prob- brown, usually glaucous or pruinose cones. Bract-
lems with identification. scale complexes 4 or 6(–8), decussate or in whorls of
IUCN: DD 3 (or 2 whorls + 2 opposite), entirely fused; bract tips
often conspicuous, up to 1 mm, on a small transverse
ridge or elevation; scale tissue dry and fibrous, with
Juniperus deppeana Steud., Nomencl. Bot., ed. 2, 1: hardening outer layer. Seeds (1–)2–4(–6) per cone
835. 1840. (usually 3), ovoid, more or less curved or flattened, 405
often unequal in size, up to 8 × 5 mm, lustrous brown
towards apex, with resinous pits and adhering scale
Etymology
tissue towards base.
This species was named after Ferdinand Deppe
(1794–1861) who collected plants in Mexico with
Distribution
Christian Julius Wilhelm Schiede in 1828–29.
SW USA: Arizona, New Mexico, W Texas; Mexico.
TDWG codes: 76 ARI 77 NWM TEX 79 MXC-PU
Vernacular names
MXE-CO MXE-CU MXE-DU MXE-HI MXE-ZA MXG-
Alligator Juniper, Checkerbark Juniper; Cedro, Ce- VC MXN-SO MXS-OA
dro chino, Táscate, Táscate blanco (Spanish); Tláscal
(Aztec)
Ecology
In grassland, scrubland or (grazed) Pinyon-Juniper
Description
woodland and open pine forests, with e.g. Opuntia,
Arborescent shrubs or trees to 12–15(–25) m, dio- Agave, Cactaceae, Yucca, and Solanum papita in open
ecious; monopodial; trunk very short or longer, to habitats; with Pinus cembroides, P. leiophylla, P. teo-
1.2(–2) m diam. Bark on trunks 1–5 cm thick (oc- cote, Juniperus flaccida, and Quercus spp. in scrub-
casionally to 20 cm at base), hard, scaly, breaking land, woodland and forest; in semi-arid mountains
up into tesselated patterns or longitudinal fissures and the more arid intermountain zone on dry, rocky
and reticulating ridges, slowly exfoliating with small slopes or in valleys and flats with sandy soils, on
flakes or long strips; inner bark red-brown or dark limestone, volcanic deposits or other types of rock.
brown; outer bark (ash-)grey. Branches long, spread- The altitudinal range for the species is from 750 m
ing or assurgent, forming a broad, rounded or py- to 2750 m a.s.l.
ramidal crown. Foliage branchlets ultimately slender,
stiff or more or less flaccid, up to 25 mm long, quad-
Uses
rangular or subterete, (0.8)1–1.4 mm wide, covered
with imbricate (gibbous) leaves, persistent. Leaves The wood may be used locally for small scale build-
on mature plants scale-like, on ultimate branch- ing and fence posts for farms, but is not exploited
lets decussate, imbricate, decurrent with appressed commercially. Larger trees have been felled for their
apex, 1.0–2.5 × 0.6–1.2 mm, ovate-rhombic, convex durable timber especially in parts of Mexico. In
or keeled; margins hyaline and minutely denticulate; horticulture Alligator juniper is suitable as an orna-
abaxial stomata in 2 conspicuous bands, adaxial sto- mental tree in regions with hot, dry summers and
mata in two concave fields; leaves abaxially glandular can withstand winter frost well. A few cultivars are
or eglandular; gland in central depression, elliptic or known and the species shows considerable variation
rounded, active with resin exudate or inactive; leaf within its natural range, which could lead to further
colour yellowish green to light green. Pollen cones selections if the effort was made. In Mexico I have
numerous, solitary, terminal on branchlets, oblong seen it (planted?) as a boundary tree between fields
to cylindrical, 4–6 × 1.6–2.1 mm; microsporophylls in the state of Puebla, together with Agave ameri-
12–16, decussate, peltate, with ­obtuse or nearly acute cana.
 5 varieties are recognized:
Conservation
Juniperus deppeana Steud. var. deppeana. Type: IUCN: LC
Mexico: Veracruz, Perote, Cofre de Perote, (Volcán
Nauhcampatepetl), C. J. W. Schiede s.n. (lectotype Juniperus deppeana Steud. var. robusta Martínez,
MO). Fig. 137, 138 Anales Inst. Biol. Univ. Nac. México 17 (1): 47.
1946. Type: Mexico: Durango, Pueblo Nuevo, Pino
Gordo, C. E. Blanco A 502 (holotype MEXU).
Description
Trees to 15 m. Bark tesselated on trunk and large Juniperus patoniana Martínez, Anales Inst. Biol.
406 branches. Foliage branchlets ultimately quadrangu- Univ. Nac. México 27: 62. 1946; Juniperus deppeana
lar. Leaves mostly eglandular but usually glandular Steud. var. patoniana (Martínez) Zanoni, Phytologia
on penultimate branchlets, with obtuse or acute 38: 438. 1978.
apex. Seed cones 7–12 mm diam., rarely ovoid and
then up to 15 mm long, reddish brown, often glau-
Description
cous but not nearly white-pruinose.
Trees to 20–25 m; bark usually tesselated only near
base of trunk, breaking in large segments to 5 cm,
Distribution
further upwards fissured, sometimes with reticulate
SW USA: Arizona, New Mexico, W Texas; Mexico. ridges, scaly, exfoliating with longitudinal strips.
TDWG codes: 76 ARI 77 NWM TEX 79 MXC-PU Leaves eglandular or with inconspicuous gland;
MXE-CO MXE-CU MXE-HI MXE-ZA MXG-VC MXS- apex obtuse or acute. Seed cones globose, large, 10–
OA MXN-SO 15 mm diam., glaucous.

Conservation Distribution
IUCN: LC Mexico: Chihuahua, Durango, Zacatecas; USA: Ari-
zona.
Juniperus deppeana Steud. var. pachyphlaea (Torr.) TDWG codes: 76 ARI 79 MXE-CU MXE-DU MXE-
Martínez, Anales Inst. Biol. Univ. Nac. México 17 ZA
(1): 53. 1946. Type: USA: New Mexico, Valencia Co.,
Zuni Mts., J. M. Bigelow s.n. (holotype NY).
Ecology
Predominantly in pine-oak woodland or pine forest,
Description
with Pinus cembroides, Quercus spp., Arctostaphylos,
Arborescent shrubs or small trees to 10 m. Bark Commelina, Dahlia, Geranium, Sedum and other
tesselated on trunk. Leaves on ultimate branchlets herbs; in mountains on rocky soil from 1000 m to
small, 1–1.8 mm long and often as wide, always glan- 2700 m a.s.l.
dular; glands often with white drying resinous exu-
date, keeled towards the obtuse or acute apex. Seed
Conservation
cones as in var. deppeana.
The total area in which this variety is known is more
limited than that for the species. It appears that this
Distribution
form is more often involved in changes in land use
USA: Arizona, New Mexico, W Texas; Mexico: So- leading to situations where scattered trees may be
nora. preserved as shade trees but chances of natural re-
TDWG codes: 76 ARI 77 NWM tex 79 MXN-SO generation seem greatly reduced.
IUCN: VU (B1+2b)
Juniperus deppeana Steud. var. sperryi Correll, 1–1.3 mm diam., subterete, rarely quadrangular.
Wrightia 3: 188. 1966. Type: USA: Texas, Jeff Davis Leaves ovate with acuminate or mucronate apex,
Co., Davis Mts., ca. 13 km from Sproul Ranch HQ, glandular. Seed cones globose, 10–15 mm diam.,
O. E. Sperry T 870 (holotype GH). light brown, strongly whitish pruinose.

Description Distribution
Arborescent shrubs or small trees. Bark fissured Mexico: Durango, W Zacatecas.
from base of trunk upwards, shaggy, slowly ex­ TDWG codes: 79 MXE-DU MXE-ZA
foliating with long strips. Foliage branches droop-
ing, ultimate branchlets more or less flaccid and 407
Conservation
pendulous. Leaves with inconspicuous, but some-
times active gland. Only known from a limited area within which it is
uncommon and in which a decline is inferred from
deforestation.
Distribution
IUCN: VU (B1+2b)
USA, W Texas, described from two trees in the
Davis Mountains and a collection made in 1931 in
the Guadalupe Mountains. One known locality in Juniperus drupacea Labill., Icon. Pl. Syriae 2: 4,
Sonora, Mexico. Other scattered trees probably exist t. 8. 1791. Arceuthos drupacea (Labill.) Antoine &
in canyons of the semiarid mountains in the region. Kotschy, Oesterr. Bot. Wochenbl. 4 (31): 249. 1854.
TDWG codes: 77 TEX 79 MXN-SO Type: Turkey: Hatay, Jebel Akra, [“Habitat in monte
Cassio quem Arabi vocant djebel lacara”], J. J. H. de
la Labillardière s.n. (holotype G-DEL?, isotypes FI,
Ecology
K). Pl. 15
Scattered trees on slopes of canyons in semi-arid
mountains. One tree grows in grass and scrub on
Etymology
rocky soil (scree) with no other trees in the vicinity.
The known altitudinal range is 1800–2000 m. The species epithet means ‘with stone fruits’ and re-
fers to the hard seed within.
Conservation
Vernacular names
Only known from a few scattered, mature or old
trees with no signs of regeneration. Syrian juniper; andys (Syria); enek (Turkish)
IUCN: CR (D)
Description
Juniperus deppeana Steud. var. zacatecensis
Martínez, Anales Inst. Biol. Univ. Nac. México 17 Trees usually 10–15(–23 in Greece, –40 in Turkey)
(1): 57. 1946. Juniperus deppeana Steud. f. zacatensis m tall, sometimes a shrub, dioecious or rarely mon-
(Martínez) R. P. Adams, Phytologia 88 (3): 229. oecious; trunk to 1–1.2 m diam. Bark on trunk with
2006. Type: Mexico: Zacatecas, Sombrerete, 10 long, fibrous strips, brown turning grey. Branches
km E of Sombrerete, M. Martínez A 503 (holotype numerous, long, spreading or assurging, the lower
MEXU). ones more or less pendulous. Foliage usually dense,
with short, spreading, ‘jointed’ ultimate branchlets,
persistent, forming a dense, pyramidal or conical,
Description
in older trees rounded crown. Leaves in alternating
Arborescent shrubs or small trees to 5–6 m, branch- whorls of 3, articulate-decurrent, spreading 60–90
ing from near the ground, branches assurgent, degrees at nodes 3–5(–10) mm apart, rigid, jointed
forming an irregular, wide crown. Bark on trunk to leaf part adnate with shoot, ovate-lanceolate,
tesselated or more or less fissured, scaly, exfoliating but shortest foliage leaves narrowly ovate (‘boat-
with irregularly shaped flakes. Branchlets slender, shaped’), (4–)10–20(–24) × 2–4 mm, straight;
408

2 5

3
6
7

6
4

Pl ate 15. Juniperus drupacea. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Cross-section of leaf.
5. Pollen cone. 6, 7. Seed cones (with cross-section showing fused seeds).
margins entire; apex acute-pungent; epistomatic, J. oxycedrus subsp. macrocarpa (in which the 3 seeds
stomata in two conspicuous broad bands, separat- remain free but lie closely appressed so as to deter-
ed by a midrib, bordered by green margins wider mine each other’s shape during growth) suggest that
than the midrib; abaxial face prominently keeled these differences are relative rather than absolute.
or ridged, green to dark green. Pollen cones axil-
lary to slightly modified leaves of 2–3 consequtive
Distribution
whorls on ultimate branchlets, 2–3 per leaf whorl,
often appearing clustered, subglobose to ovoid, 4–7 Greece, Turkey (mountains along Mediterranean
× 4–5 mm; microsporophylls in (3–)5–6 alternating coast), Syria, Lebanon, Israel.
whorls of 3, subpeltate (leaf-like), with entire mar- TDWG codes: 13 GRC 34 LBS-LB LBS-SY PAL-IS
gins and acuminate apex, bearing 4–6(–9) abaxial TUR 409
ellipsoid pollen sacs in 2 rows near the lower margin.
Seed cones axillary on 5–10 mm long dwarf shoots
Ecology
with whorls of small (2–4 mm) triangular leaves,
maturing in 2 years; mature cones ovoid-globose, Juniperus drupacea occurs in mixed (low) montane
15–30 × 12–24 mm, with 2–3(–4) alternating, imbri- conifer forest and woodland with Pinus brutia, P. ni-
cate whorls of 3 mostly fused bract-scale complexes, gra, Abies cephalonica, A. cilicica, Cedrus libani, Ju-
(2–)3 whorls fully developed (middle scales largest), niperus excelsa, J. foetidissima, J. oxycedrus, Quercus
ovate-mucronate, thick, with prominent margins, coccifera, Q. ilex, and more rarely with Fagus orien-
becoming hard semi-woody, more or less resinous, talis. Forest degradation has changed the vegeta-
surface rugose, light to dark brown, often glaucous tion in some areas to maquis in which the junipers
or pruinose. Seeds usually 3 per cone, more or less have regenerated or spread. It occupies, with other
triangular, often with small resin vescicles on outer junipers, the rockier, more shallow soils, often on
surface, fused for ⅔-¾ proximally into an ovoid- calcareous but also on granitic rock. The altitudinal
globose yellowish brown ‘stone’ 10–18 × 7–15 mm, range is from 600 m to 1800 m a.s.l. The climate is
connate distally. Mediterranean, with winter rain, sometimes snow,
and dry, hot summers.
Taxonomic notes
Conservation
This species was known in pre-Linnaean times to
botanists as Juniperus major on account of its large This species is rare in Greece, Lebanon and Israel,
cones and tree size and botanical travellers reported where it is probably Vulnerable (Greece) or even
it from the Turkish mountains as early as 1547; de- Endangered (Israel); however in Turkey and Syria it
spite this Linnaeus (1753) appears to have overlooked is still more widespread and locally common. Bo-
it. De Labillardière encountered it in N Syria near ratynski & Browicz (1983) gave an account of the
the Turkish border on Mt. Cassius (Djebel Akra, populations in Greece, all in the Parnon Massif in
now in Turkey) in 1788. Its very large cones which the Peloponnisos. While some stands represent de-
clearly show the outline of the bract-scale complexes graded fir forest (Abies cephalonica) or have been
and contain partly fused, large seeds have been seen converted to pasture, others appear to be natural or
as sufficiently distinct to erect a new genus Arceu- semi-natural. They suggested protective status for at
thos for it. More commonly, this species has been least one of the major stands in Greece.
assigned to its own section Caryocedrus since Endli- IUCN: LC
cher (1847), although Gaussen (1968) used subgenus
Caryocedrus for it. The partly fused seeds stressed
Uses
by Endlicher as a diagnostic character of this sec-
tion (= J. drupacea), as well as the cone itself, require The wood of large trees is valuable as timber with de-
a careful ontogenetic study for which unfortunately cay resistant properties but it is not extensively used
no material was available. What is clear is that the for this purpose. In Turkey, the local population
ovules originate independently in the axils of the harvest the fleshy cones, which have a high content
fertile whorl of 3 bracts, so that fusion is a secondary of sugars, for consumption as a kind of marmelade
character state. Some similarities with large cones of or dried fruit. Its habit in cultivation tends to remain
 columnar for many years and its large leaves with × 2 mm; microsporophylls 10–12, decussate, peltate,
conspicuous stomatal bands are also ornamental. as broad as long, with hyaline-denticulate margins,
Yet its use in gardens is limited despite good growth obtuse, bearing 2–3 abaxial, relatively large pollen
rates and tolerance to frost; this omission deserves sacs. Seed cones numerous, terminal on curved or
to be rectified as it is probably a better species for straight, 2–3 mm long branchlets; mature cones sub-
horticulture than the more common J. oxycedrus globose to nearly reniform, often wider than long,
(in southern Europe and regions with equivalent 4–6 × 5–7 mm, irregular, more or less bilobed if with
­climate). 2 seeds, smooth (rugose in sicco), orange-brown or
reddish brown, often glaucous to pruinose. Bract-
scale complexes 4, decussate, upper pair larger than
410 Juniperus durangensis Martínez, Anales Inst. Biol. lower pair, bract tips exserted, 0.5 mm, triangular
Univ. Nac. México 17 (1): 94. 1946. Type: Mexico: and more or less curved; tissue soft pulpy, succulent
Durango, El Salto, Puerto de Santo Domingo, or sometimes dry. Seeds 1–3(–4) per cone, variable
M. Martínez 7015 (holotype MEXU). but mostly angular-ovoid, 3–4.5 × 2–3 mm, curved
or not, ridged and/or shallowly pitted, hilum a third
to half of seed, resinous or not, seed coat light brown
Etymology
or reddish brown, but hilum lighter and usually
The species epithet refers to the State of Durango in turning yellowish brown.
Mexico.
Distribution
Vernacular names
Mexico: Aguascalientes, Chihuahua, Durango, Jalis-
No common names were recorded for this species. co, Sonora, Zacatecas.
TDWG codes: 79 MXE-AG MXE-CU MXE-DU MXE-
ZA MXN-SO MXS-JA
Description
Shrubs or small trees to 6 m, presumably dioecious;
Ecology
trunk forked at base or branching low, or erect and
curved or contorted, to 30–40 cm diam. Bark be- Martínez (op. cit.) described the habitat of this spe-
coming fibrous, 5–10 mm thick, exfoliating in long, cies as very poor, dry, rocky soil, and presented some
thin strips, light brown weathering grey. Branches photographs of trees that are apparently growing in
thick, contorted, spreading and assurgent, foliage a deforested landscape, which probably had been
dense, forming an irregular and open crown with covered with pine-oak woodland. On two labels of
wide spreading branches. Foliage branches numer- herbarium collections examined, a habitat consist-
ous, rigid, forming dense tufts, ultimate branchlets ing of (cleared) oak-juniper woodland was men-
short, rigid, spreading in all directions, straight or tioned, with Juniperus deppeana var. robusta also
towards ends of foliage branches down-curved, present. These collections were made at altitudes of
4–10(–18) mm long, 1.2–1.5 mm diam., more or less 2500–2700 m a.s.l. More work is needed to estab-
quadrangular in cross-section, leading ultimate lish both the extent and ecology of this species in
branchlets to 2 mm diam. Leaves decussate on ulti- N Mexico.
mate lateral branchlets, in alternating whorls of 3 on
lower branchlets, imbricate, on ultimate branchlets
Conservation
appressed, broadly ovoid-rhombic, 0.8–1.2 × 0.8–1
mm, often nearly as wide as long, gibbous; margins Although it is known from a wide range in N Mex-
thick, minutely denticulate; apex obtuse, leaves on ico, probably less than 15 scattered collections have
older branchlets broad rhombic, acute; stomata few been made and it is apparently not common. Often,
on lower margin abaxially and scattered stomata these collections were made in ‘clearings’ of pine-
from base to apex adaxially; gland central, small, oak forest or woodland, with all but a few trees cut.
oval-oblong to linear, inactive; leaf colour dark It is therefore assumed that this species qualifies as
green, new foliage often yellowish green. Pollen Vulnerable, but a wider survey is desirable to estab-
cones terminal, solitary, subglobose or ovoid, 2.5–3 lish its distribution more exactly. Difficulties with
identification, as other, similar species occur in the cular, often with exudate; leaf colour yellowish green
region in similar habitat, mean that only special- to dull light green. Pollen cones numerous, solitary,
ized people are qualified to do this; its low value as terminal on short branchlets, globose to subglobose,
a timber tree provides little incentive for foresters to 3–4 × 2–3 mm; microsporophylls 8–10, decussate,
­undertake such a project. peltate, with rounded entire margins, bearing 3–4
IUCN: VU (B1+2b) abaxial pollen sacs. Seed cones terminal on short,
erect branchlets, maturing in the second season and
becoming (sub)globose, 6–12(–14) mm diam., pink-
Uses
ish brown, blue or purplish blue, usually glaucous
No specific uses have been recorded of this species; or pruinose. Bract-scale complexes 4–6, decussate,
it presumably serves both as firewood and for fence- entirely fused, sometimes sutures faintly visible near 411
posts locally. apex of cone, bract tip small, ca. 0.5 mm, on a small
ridge, scale tissue dry pulpy or rarely more suc-
culent, resinous. Seeds (2–)3–6(–8) per cone, sub-
Juniperus excelsa M.-Bieb., Tabl. Prov. Mer. ovoid-conical, more or less flattened or curved, 4–6
Casp.: 204. 1798. [Beschr. Länd. Terek & Kur, Bot. × 3–4 mm, yellowish to reddish brown, with lighter
Anhang: 204. 1800.] hilum at base.

Etymology Distribution
The species epithet means ‘high’ and perhaps refers SE Europe: Albania, Bulgaria, Greece, Macedonia;
to the considerable size of this species compared to E Europe: Ukraine: Krym [Crimea]; Caucasus: Ar-
the shrubby species known from central and west- menia, Azerbaijan, Gruzija, Russia(?); Central Asia:
ern Europe. Kazakhstan, Kirgyzstan, Tadzhikistan, Turkmeni-
stan, Uzbekistan; W Asia: Afghanistan, Cyprus,
Iran, Lebanon, Syria, Turkey; Arabian Peninsula:
Vernacular names
Oman; S. Asia: NW India, Pakistan.
Greek juniper, Grecian juniper, Crimean juni- TDWG codes: 13 ALB BUL GRC YUG-MA 14 KRY
per, Turkestan juniper (subsp. polycarpos); Ardich 32 KAZ KGZ TKM TZK UZB 33 TCS 34 AFG CYP IRN
(Turkish); Dedali-gwia (Iran); Parmiro alañ (Cen- LBS-LB LBS-SY TUR 35 OMA 40 PAK WHM-JK
tral Asia, subsp. polycarpos)
Ecology
Description
In shrubland (Mediterranean), deciduous wood-
Trees (rarely shrubs) to 20–25 m, monoecious or land and coniferous forest, in the interior eastern
more commonly dioecious; monopodial; trunk up parts of its range also in montane Seriphidium mar-
to 2.5 m d.b.h., or a (decumbent) multistemmed itimum (Artemisia maritima) steppes and on alpine
shrub. Bark on trunks exfoliating in longitudinal fi- scree or moraines. The altitudinal range is (100–)
brous strips, reddish brown weathering grey-brown. 500–3000(–3950) m a.s.l. It is most common on
Branches spreading or ascending, foliage branches dry slopes of calcareous hills and mountains in the
very dense, short, stiff and spreading, or longer and Mediterranean climate parts of its range, but occurs
more or less lax, forming a dense, rounded or irreg- also on siliceous rock in the continental interior,
ular, sometimes sympodial crown. Foliage branch- more often near intermittent groundwater sources.
lets ultimately subterete or weakly quadrangular, In Central Asia, it is restricted to the more west-
0.7–1.3 mm wide, covered with closely appressed ern parts of the numerous mountain ranges, where
leaves, persistent. Leaves on mature plants normally it forms, with some other species, (J. semiglobosa,
scale-like, decussate, imbricate, decurrent, 0.6–1.6 J. pseudosabina, J. sabina) extensive juniper forest
× 0.4–0.9 mm, ovate-rhombic, obtuse; margins en- or woodland (‘Archa forest’) on all but the N-facing
tire; stomata in 2 inconspicuous lines on each side slopes (where e.g. Picea schrenkiana dominates, a
mostly near base; leaves abaxially glandular; gland conifer that is more abundant and not restricted to
central, large and conspicuous, elliptic or nearly cir- N-facing slopes farther east). In contrast to North
 America, this juniper forest is not mixed with pines
Ecology
(Pinus spp.) and angiosperms are uncommon in it.
Several species of Juniperus, including low shrubs, This subspecies occurs in shrubland (Mediterra-
therefore determine the structure of these forests or nean), woodland and pine forest on dry hillsides
woodlands in large areas, often from the valley bot- and rocky mountain slopes at altitudes from 100 m
tom to the tree line. to 2000(–2700) m a.s.l. It is associated with Pinus
brutia, P. nigra, Cedrus libani, Abies cephalonica,
A. cilicica, A. nordmanniana, Juniperus foetidissima,
Uses
J. oxycedrus, Quercus spp., Pistacio lentiscus, Ber-
The wood of J. excelsa is durable and hard and, in beris, Acer obtusifolium, etc. It grows most often
412 parts of Turkey at least, sufficiently common to be on limestone and also on igneous rocks, but not in
utilized. However, trees grow slowly and take two acidic soil. The climate is Mediterranean, with win-
centuries or more to attain any harvestable size, so ter rain and dry, hot summers, grading eastward
its exploitation for timber is or would not be sus- into more continental conditions, where J. excelsa
tainable if it occurred on a commercial scale. In subsp. polycarpos predominates.
many parts of SW Asia it also serves as firewood,
especially for people who live (temporarily) in the
Conservation
mountains. The wood of large trees is used for car-
pentry and furniture. Foliage is sold and burnt as IUCN: LC
incense. Greek juniper is uncommon in horticul-
ture and only a few cultivars have been selected; the Juniperus excelsa M.-Bieb. subsp. polycarpos
one best known in Britain is a columnar form with (K. Koch) Takht., Fl. Yerev.: 53. 1972. Juniperus
glaucous and more juvenile type foliage properly polycarpos K. Koch, Linnaea 22: 303. 1849; Juni-
named ‘Stricta’ but also known under several vari- perus excelsa M.-Bieb. var. polycarpos (K. Koch)
etal names. Forms with pendulous foliage and with Silba, Phytologia Mem. 7: 34. 1984. Type: Turkey:
dwarf growth have also been named and are used Gümüshane, Anadolu Daglari, Taltaban, P. E. E. Sin-
more often in central and southern Europe. tenis 5520 (neotype L). Pl. 16

2 subspecies are recognized: Juniperus macropoda Boiss., Fl. Orient. 5: 709. 1884.
Juniperus seravschanica Kom., Bot. Zurn. (Moscow
Juniperus excelsa M.-Bieb. subsp. excelsa. Type: & Leningrad) 17: 481. 1932; Juniperus poly­carpos
Ukraine: Crimea, Krymskiye Gory, P. S. Pallas, [ex K. Koch var. seravschanica (Kom.) Kitam., [Fl. Pl.
herb. Pallas] s.n. (lectotype LE). W. Pakist. Afghan.: 7. 1964] Add. & Corr. Fl. Afghan.:
68. 1966; Juniperus excelsa M.-Bieb. subsp. serav­
schanica (Kom.) Imkhan., Bot. Zurn. 75 (3): 407. 1990.
Description
Juniperus turcomanica B. Fedtsch., in Fedtschenko
Trees to 20–25 m; ultimate branchlets slender, 0.7–1 et al., Fl. Turkmenii 1: 14. 1932; Juniperus excelsa M.-
mm diam., weakly quadrangular or subterete, often Bieb. subsp. turcomanica (B. Fedtsch.) Imkhan., Bot.
in regularly disposed sprays, more or less lax. Seed Zurn. 75 (3): 408. 1990; Juniperus polycarpos K. Koch
cones 6–11 mm diam., mostly darker hues, glaucous var. turcomanica (B. Fedtsch.) R. P. Adams, Phytolo-
or pruinose. Seeds 3–6(–8) per cone. gia 86 (2): 50. 2004.
Juniperus polycarpos K. Koch var. pendula Mulk.,
Dokl. A. N. Armen. SSR 45 (2): 86. 1967; Juniperus
Distribution
excelsa M.-Bieb. subsp. polycarpos (K. Koch) Takht.
SE Europe: Albania, Bulgaria, Greece, Macedonia; var. pendula (Mulk.) Imkhan., Bot. Zurn. 75 (3): 407.
E Europe: Ukraine: Krym [Crimea]; Caucasus: Ar- 1990.
menia, Azerbaijan, Gruzija; W Asia: Cyprus, Iran
(Elburz Mts.), Lebanon, Syria, Turkey; Central Asia:
Description
Turkmenistan (Kopet Mts.).
TDWG codes: 13 ALB BUL GRC YUG-MA 14 KRY 32 Trees to 20 m, rarely (decumbent) shrubs; ultimate
TKM 33 TCS 34 CYP IRN LBS-LB LBS-SY TUR branchlets thicker (1–1.3 mm diam.), subterete, ­rigid,
 413

6
1
8

10

2 9
3

Pl ate 16 . Juniperus excelsa subsp. polycarpos. 1. Habit of tree. 2. Branch with foliage and seed cones.
3. Branchlet with adult leaves. 4. Branchlet with juvenile leaves. 5. Seedling. 6, 7. Pollen cones. 8. Branchlet
with seed cones. 9. Seed cone. 10. Opened seed cone with seeds.
 spreading in all directions. Seed cones 8–12(–14)
Description
mm diam., pinkish brown to purplish blue, usually
pruinose. Seeds (2–)3–4(–6) per cone. Small trees (rarely shrubs) to 10–15 m, dioecious;
trunk very short to 3 m, to 50–80 cm diam. Bark
on trunks ca. 10 mm thick, fissured, scaly, exfo-
Distribution
liating in fibrous strips, brown weathering grey-
Central Asia: Kazakhstan, Kirgyzstan, Tadjikistan, brown. Branches long, spreading or ascending,
Turkmenistan, Uzbekistan; Caucasus: Armenia, Az- foliage branches long and lax, spreading to pendu-
erbaijan, Russia (?); W Asia: Afghanistan, Iran, E lous, commonly in more or less flattened sprays,
Turkey; Arabian Peninsula: Oman; S Asia: Pakistan, forming a sympodial, broad, rounded crown. Foli-
414 NW India (Kashmir). age branchlets ultimately slender, flaccid, subterete,
TDWG codes: 32 KAZ KGZ TKM TZK UZB 33 TCS 34 1–1.2 mm wide, covered with imbricate leaves, per-
AFG IRN TUR 35 OMA 40 PAK WHM-JK sistent. Leaves on mature plants scale-like, decus-
sate, imbricate, decurrent with free to spreading
apex, 2–4 × 0.7–1 mm, broad lanceolate to rhom-
Ecology
bic, acute to acuminate-pungent; gland abaxially
The continental subspecies occupies a range of in central depression, elliptic or oblong; margins
habitats from montane (mixed) coniferous forest to nearly entire-hyaline or minutely denticulate; leaf
upper montane steppes dominated by ­Seriphidium colour yellowish green to light green; stomata abaxi-
maritimum (Artemisa maritima). Its altitudinal ally in 2 inconspicuous lines near the base, adaxially
range is (550–)1000–3300(–3950) m a.s.l. In coni­ in two bands barely separated by a midrib. Pollen
ferous forest it can be associated with Pinus gerardi- cones numerous, solitary, terminal on short branch-
ana, P. wallichiana, Cedrus deodara or Abies pindrow, lets, ovoid-oblong, 3–5 × 1.5–2 mm, often elongated
in broad-leaved forest or woodland with Carpinus at pollen dispersal; microsporophylls 10–16, decus-
sp., Juglans nigra, Malus sieversii, Crataegus sp., Pru- sate, peltate-acuminate, with hyaline-erose margins,
nus sp., Sorbus sp. and shrubs e.g. Dodonaea and bearing 2–4 abaxial pollen sacs. Seed cones terminal
Sageretia. At its highest altitudes in Afghanistan and on very short, erect lateral branchlets, maturing in
Pakistan it forms very open stands on bare scree or the second season to (sub)globose, 8.5–16 (in one
glacial moraines with decumbent Juniperus squa- var. up to 20, in another down to 6) mm diam., light
mata. It is found on calcareous as well as siliceous brown, usually glaucous or pruinose cones. Bract-
rocks, usually in shallow, stony soils, but sometimes scale complexes 6, decussate, entirely fused, sutures
in loess or loam, and on dry slopes or near intermit- often visible on entire cone forming a polygonal pat-
tent streams. tern; bract tip usually conspicuous, triangular, ca.
1 mm, on a small transverse ridge; scale tissue dry
pulpy. Seeds (4–)6–8(–12) per cone, angular-conical,
Conservation
more or less flattened, 3–4 mm long, reddish brown,
IUCN: LC with lighter hilum at base.

Distribution
Juniperus flaccida Schltdl., Linnaea 12: 495. 1838.
SW USA: W Texas (Big Bend National Park); Mex-
ico: Chihuahua, Coahuila, Durango, Guerrero,
Etymology
Hidalgo, Jalisco, México, México D. F., Michoacan,
The species epithet (Latin flaccidus = soft, weak, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro,
withered) refers to the pendulous foliage branchlets. San Luis Potosí, Sonora, Tamaulipas, Tlascala, Ver-
acruz, Zacatecas.
TDWG codes: 77 TEX 79 MXC-DF MXC-ME MXC-
Vernacular names
MO MXC-PU MXC-TL MXE-CO MXE-CU MXE-DU
Drooping Juniper, Weeping Juniper, Mexican MXE-GU MXE-HI MXE-NL MXE-QU MXE-SL MXE-
Drooping Juniper; Cedro liso, Enebro, Tascate TA MXE-ZA MXG-VC MXN-SO MXS-GR MXS-JA
(Spanish); Tláscal (Aztec) MXS-MI MXS-OA
 Juniperus flaccida Schltdl. var. martinezii (Pérez
Ecology
de la Rosa) Silba, Phytologia 58: 367. 1985. Juniperus
Juniperus flaccida occurs in montane Pinyon-Juni- martinezii Pérez de la Rosa, Phytologia 57: 81.
per woodland, pine-oak forest and woodland, and 1985. Type: Mexico: Jalisco, Sierra Cuatralba, El
oak woodland, with Pinus cembroides, Pinus spp., Cuarento, 4 km E of village, J. A. Pérez de la Rosa
Juniperus deppeana, J. saltillensis, Quercus spp., 661 (holotype IBUG).
Fraxinus, Arbutus, mimosoid legumes, Yucca spp.,
etc., on limestone or other rocks including siliceous
Description
(granitic) rocks. The altitudinal range is from 800 m
to 2600 m a.s.l. Leaves with hyaline, (minutely) denticulate margins,
acuminate-pungent, glaucous green. Seed cones 415
small, 6–8 mm diam., with only 1–3 seeds.
Uses
No uses have been recorded for this species; the
Distribution
wood may locally serve for fuel or be used for fence
posts. Mexico: Jalisco (Ojuelos, Sierra Cuatralba).
TDWG codes: 79 MXS-JA
3 varieties are recognized:
Conservation
Juniperus flaccida Schltdl. var. flaccida. Type:
Mexico: Hidalgo, Regla, C. A. Ehrenberg 8 Only known from a limited area in which forest
(lectotype MO). Fig. 139 ­cover is decreasing overall, this variety could be at
some risk of extinction.
IUCN: NT
Description
Foliage spreading to pendulous, not in more or Juniperus flaccida Schltdl. var. poblana Martínez,
less flattened, distichously branching sprays, green. Anales Inst. Biol. Univ. Nac. México 17 (1): 31.
Leaves with nearly entire-hyaline margins, acumi- 1946. Juniperus poblana (Martínez) R. P. Adams,
nate-pungent. Seed cones 8.5–15 mm diam., with Phytologia 88 (3): 239. 2006. Type: Mexico: Puebla,
more than 3 and up to 12 (usually 6–8) seeds. Amozoc de Mota, 1 km NW of town, M. Martínez
A 507 (holotype MEXU).
Distribution
Description
SW USA: W Texas (Big Bend National Park); Mex-
ico: Chihuahua, Coahuila, Durango, Guerrero, Foliage in more or less distichously branched, slight-
Hidalgo, Jalisco, México, México D. F., Michoacan, ly flattened sprays; leaves acute. Seed cones large, up
Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, to 20 mm diam., with conspicuous sutures of bract-
San Luis Potosí, Sonora, Tamaulipas, Tlascala, Ver- scale complexes (polygonal markings).
acruz, Zacatecas.
TDWG codes: 77 TEX 79 MXC-DF MXC-ME MXC-
Taxonomic notes
MO MXC-PU MXC-TL MXE-CO MXE-CU MXE-DU
MXE-GU MXE-HI MXE-NL MXE-QU MXE-SL MXE- In a recent paper, Adams et al. (2006) concluded
TA MXE-ZA MXG-VC MXN-SO MXS-GR MXS-JA that this variety is distinct based on molecular data
MXS-MI MXS-OA (DNA sequence data and RAPDs) and should be
recognized as a species. Only one sample of this
taxon was included in the analysis and its phyloge-
Conservation
netic relationship remained unresolved although it
IUCN: LC seemed distant from J. flaccida var. flaccida and var.
martinezii.
 crushed. Pollen cones numerous, solitary, terminal
Distribution
on short branchlets, subglobose to ovoid-globose,
Mexico: Coahuila, Guerrero, Jalisco, Michoacan, 2–3.5 mm long; microsporophylls 8–12, decussate,
Nuevo León, Oaxaca, Puebla, Zacatecas. peltate, with rounded entire or erose margins, bear-
TDWG codes: 79 MXC-PU MXE-CO MXE-NL MXS- ing 3–4 abaxial pollen sacs. Seed cones terminal on
GR MXS-JA MXS-MI MXS-OA 3–4 mm long, erect branchlets, maturing in the sec-
ond season to become (sub)globose, 5–13 mm diam.,
dark blue or blackish blue, usually glaucous or pru-
Conservation
inose. Bract-scale complexes 6, decussate, entirely
IUCN: LC fused, sometimes sutures visible, bract tip small, ca.
416 0.5 mm, on a small ridge, smooth or more or less
rugose in sicco; scale tissue dry pulpy or rarely more
Juniperus foetidissima Willd., Sp. Pl. 4 (2): 853. succulent, resinous. Seeds 1–2(–3) per cone, if more
1806. Type: Turkey: locality unknown [“habitat in than one, connate, often partly fused, ovoid-globose
Armenia”], J. P. de Tournefort B-W 18547 (holotype or more or less flattened on one side or hemispheri-
B-W). cal, 5–7 mm diam., pale brown.

Etymology Distribution
The species epithet means ‘stinking extremely’ and SE Europe: Albania, Greece, Macedonia; E Europe:
refers to the odour of crushed foliage. Ukraine: Krym [Crimea]; Caucasus: Armenia,
­Azerbaijan, Gruziya, Krasnodar, Stavropol; W Asia:
Cyprus, NW Iran, Lebanon, Turkey; Central Asia:
Vernacular names
Turkmenistan.
Stinking juniper; Selwi Aghatch, Ardyoch, Selwi Ar- TDWG codes: 13 ALB GRC YUG-MA 14 KRY 32 TKM
diç (Turkey); Twia, Malokedra Mamaligwia (Greece 33 NCS-KR NCS-ST TCS-AR TCS-AZ TCS-GR 34 CYP
& Macedonia) IRN LBS-LB TUR

Description Ecology
Trees (rarely shrubs) to 25(–42) m, monoecious or In (Mediteranean) scrubland, Juniperus-Quercus
dioecious; trunk up to 3.5(–4.6) m d.b.h., or a (de- scrub, open pine forest and other coniferous for-
cumbent) multistemmed shrub. Bark on larger est, with Pinus brutia, P. nigra, Juniperus excel-
stems exfoliating in longitudinal fibrous strips, sa, J. drupacea, J. oxycedrus, Cedrus libani, Abies
weathering grey-brown. Branches spreading or as- ­cephalonica, A. cilicica, A. nordmanniana, Quercus
cending, foliage branches short, stiff and spread- ilex, Pistacia lentiscus, etc. Often on limestone crags
ing, forming a dense, broad pyramidal to eventually or rocky slopes, or on serpentine or other base-rich
rounded or irregular crown. Foliage branchlets ulti- rock, in shallow soils. The altitudinal range is from
mately subterete or weakly quadrangular, 1.2–2 mm 700 m to 2000 m a.s.l. The climate is Mediterranean
wide, covered with appressed or apically spreading with transitions to more continental interior regions
leaves, persistent. Leaves on mature plants eventu- where winters are drier and colder.
ally scale-like, decussate, imbricate, decurrent, 2–3
× 1–1.5 mm, ovate-rhombic, more or less acute; mar-
Conservation
gins entire; stomata in 2 inconspicuous lines on each
side mostly near the base; scale leaves abaxially glan- This widespread species is not threatened with ex-
dular, with central, inconspicuous gland; leaf colour tinction. Some exceptionally large and old trees, e.g.
yellowish green to lustrous green. Intermediate leaf in southern Turkey, Mersin, Kadincik Deresi, are
shapes, as well as juvenile-type leaves after coppic- protected.
ing, usually present in mature trees but diminish- IUCN: LC
ing in older trees. Foliage with a ‘foetid’ odour when
 ­ idrib, bordered by green or glaucous green mar-
m
Uses
gins as wide as midrib. Pollen cones axillary, solitary,
Large trees have been utilised in the past for the 1–2 per leaf whorl, subglobose to ovoid, 4–6 × 3–4
making of durables like furniture, but its commer- mm; microsporophylls 9–12, in alternating whorls of
cial use is now limited. In some areas firewood may 3, peltate, with hyaline margins and acuminate apex,
still be extracted from this species. It is uncommon bearing 4–6 abaxial pollen sacs near lower margin.
in cultivation, perhaps due to the unpleasant smell Seed cones axillary on short (1–2 mm) dwarf shoots
of the foliage, although this is not as extreme as its with whorls of minute scale leaves, maturing in 1
species name implies and will usually only be no- year; mature cones (sub)globose, 6–9 mm diam.,
ticed when fresh leaves are crushed. with 2 whorls of 3 completely fused bract-scale com-
plexes, only upper whorl fully developed, with 3 su- 417
tures visible on distal part of cone; bract tip hidden
Juniperus formosana Hayata, J. Coll. Sci. Imp. or very small; tissue dry pulpy, occasionally more
Univ. Tokyo 25 (19): 209. 1908. Type: Taiwan: succulent; mature cones orange-brown or light
Nantou, Chia-i Pref., Yu-Shan, [“Mt. Morrison”], brown, sometimes dark purple, slightly pruinose.
T. Kawakami & U. Mori 2039 (lectotype TI). Seeds 2–3 per cone, subglobose to (angular-)ovoid,
4–5 × 3–3.5 mm, light brown, with 3–4 basal resin
Juniperus mairei Lemée & Lév., Monde Pl. 2 (16): 20. pits and obtuse apex.
1914; Juniperus formosana Hayata var. mairei (Lemée
& Lév.) R. P. Adams & C. F. Hsieh, Biochem. Syst.
Distribution
Ecol. 30: 239. 2002.
China: S Anhui, Chongqing, W Fujian, E Gansu,
Guizhou, W Hubei, S Hunan, S Jiangsu, Jiangxi,
Etymology
NE Qinghai, S Shaanxi, Sichuan, S Xizang [Tibet],
The species epithet indicates the island of Taiwan Yunnan, Zhejiang; Taiwan.
(formerly Formosa), from where it was first de- TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
scribed; see also Latin formosus = handsome or well SC CHC-YN CHN-GS CHN-SX CHQ CHS-AH CHS-FJ
formed. CHS-HN CHS-JS CHS-JX CHS-ZJ CHT 38 TAI

Vernacular names Ecology

Formosan juniper, Prickly cypress; ci bai (Chinese) In coniferous forest, secondary coniferous wood-
land, and (subalpine) grassland, at high altitudes
predominantly on S-SW facing slopes with shrubs
Description
and grasses, also in open areas of deciduous broad-
Shrub or tree to 15(–25) m, dioecious; often leaved forest on dry, rocky soils. The altitudinal
2–3-stemmed or low branching. Bark on trunks with range is from 400 m to 3830 m a.s.l. This species
long, thin strips, grey-brown. Branches spreading or is associated with Pinus spp. Picea spp., Abies spp.,
ascending. Foliage dense or more open, mostly with Tsuga spp., Juniperus squamata, Rhododendron spp.,
pendulous, ‘jointed’ ultimate branchlets, but spread- numerous angiosperm trees and shrubs, and with
ing and rigid at high-altitude, persistent, forming grasses. It occurs on all types of rock, from lime-
a pyramidal to conical crown, or a dense shrub at stone and shale to granite.
high altitude. Leaves in alternating whorls of 3, non-
decurrent, spreading 30–60(–90) degrees at nodes
Conservation
3–10 mm apart, rigid, jointed to leaf part adnate with
triangular shoot, linear-lanceolate, (4–)10–20(–30) IUCN: LC
× (1–)1.1–2.1 mm, straight or slightly curved, usu-
ally broadest near the middle; abaxial face (weakly)
Uses
keeled, lustrous green or dark green; margins en-
tire; apex acuminate or acute-pungent; epistomatic, Formosan juniper is in cultivation and usually grows
stomata in two conspicuous bands separated by a into a columnar habit when planted in gardens and
 parks. It is more commonly in use as an ornamental abaxial pollen sacs. Seed cones terminal, matur-
in China and Japan than in Europe and the USA, ing in one year to globose, subglobose or irregular
probably because of the availability of similar forms shapes (4–)6–8(–9) mm diam., ripening to pinkish
or cultivars of J. communis there. or reddish brown, sometimes glaucous. Bract-scale
complexes 4, decussate, entirely fused and invis-
ible in mature cone, hard, dry and resinous; bract
Juniperus gamboana Martínez, Anales Inst. Biol. tip minute but visible. Seeds 1(–2) per cone, broadly
Univ. Nac. México 15 (1): 7. 1944. Juniperus deppea- ovoid or ovoid-conical, 6–7 × 5–6 mm, slightly flat-
na Steud. var. gamboana (Martínez) R. P. Adams, tened, with deep grooves and resinous pits, yellow-
Phytologia 88 (3): 229. 2006. Type: Mexico: Chia- ish brown, ochraceous or light chestnut-brown.
418 pas, Teopisca, [protologue: “inter Rancho Mitzitón
et Comitán”; photo caption: “Belén”], M. Martínez
Taxonomic notes
6701 (holotype MEXU).
This species was recently found to be closely re-
lated to Jupiperus deppeana in an analysis of DNA
Etymology
sequence data (Adams & Schwarzbach, 2006) and
This species was named after Rafael P. Gamboa, at therefore interpreted as a variety of that species.
the time Governor of the State of Chiapas, Mexico. While it shares some characters, like the checkered
bark (not present in all varieties of J. deppeana), it
also has some distinct ones. The analysis presented
Vernacular names
is far from convincing, with largely unresolved rela-
No common names have been recorded for this spe- tionships among the varieties of J. deppeana, but high
cies. statistical support for a clade with J. gamboana and
J. deppeana var. robusta. If J. gamboana were indeed
most closely related to J. deppeana var robusta, the
Description
geographical distribution of these two taxa would be
Shrubs or small trees to 8–12 m, monoecious or hard to explain, as they are separated by populations
dioecious; trunk very short or straight, to 50–90 of J. deppeana var. deppeana. A better resolved clado-
cm diam. Bark on trunks fissured, 10–15 mm thick gram based on a wider sampling of data is needed to
and hard, forming longitudinal plates, lower part justify this new taxonomy and J. gamboana is there-
of trunk often with quadrangular plates, weather- fore here maintained as a distinct species.
ing grey. Branches spreading or ascending, forming
a broadly conical or pyramidal crown, in old trees
Distribution
becoming irregular and spreading. Foliage branches
spreading or drooping, ultimate branchlets spread- Mexico: Chiapas; Guatemala: Huehuetenango.
ing at 30–50°, variable in length from 4–20 mm, TDWG codes: 79 MXT-CI 80 GUA
1–1.2(–1.4) mm thick, quadrangular to terete, per-
sistent. Leaves on ultimate branchlets decussate, on
Ecology
older branchlets often in alternate whorls of 3, or a
mixture of these arrangements, all scale-like, imbri- This species occurs in the understorey of, or mixed
cate, appressed, broadly rhombic, (1.3)1.5–2 × 0.7–1 with other trees in Quercus-Pinus-Juniperus forests,
mm (on older branchlets to 2.5 mm long), gibbous bearing many epiphytes indicating frequent fog con-
or keeled towards the acute-mucronate apex; mar- ditions, on often rocky soil types; also in open pine
gins minutely denticulate; apex more or less free esp. forest, e.g. with Pinus oocarpa, where J. gamboana
on whip shoots with acuminate leaves; stomata few attains greatest size. The known altitudinal range is
near abaxial base, in two narrow converging bands 1820–2200 m a.s.l.
adaxially; glands in a central abaxial groove, not
producing a resin drop (inactive); leaf colour light
Conservation
yellowish green. Pollen cones terminal, solitary, ob-
long, to 6 × 2 mm; microsporophylls 10–12, decus- Due to its very limited range (most collections
sate, subpeltate, with mucronate apex, bearing 3–4 known are from Chiapas, Mexico) in an area which
suffers rapid deforestation, the species is likely to invisible. Seeds 1–2 per cone, ovoid-globose, 2–3
have suffered a significant reduction in its area of mm diam., light brown.
occupancy (AOO) as well as its overall numbers of
mature trees. The situation in Guatemala with this
Distribution
species is poorly known.
IUCN: VU (B1+2b) Hispaniola (Dominican Republic and Haiti), very
localized in relict populations.
TDWG codes: 81 DOM HAI-HA
Juniperus gracilior Pilg., in Urban, Symb. Antill. 7:
481. 1913.
Ecology
419
The three varieties each occur in very limited locali-
Etymology
ties in the humid montane forest zone from 1200 m
The epithet (Latin gracilis = slender) means ‘com- to 2550 m a.s.l.; their habitat is characterized by a
paratively slender’. scattering of Manacla palms and abundant epihytes.
The forest cover has been destroyed in most locali-
ties and secondary vegetation predominates, often
Vernacular names
dominated by the fern Pteridium aquilinum. In most
Sabina, Cèdre (Spanish/French) localities the junipers are present with only solitary
individuals. The species is mainly found on acid for-
est soil, but J. gracilior var. urbaniana occupies a very
Description
limited area near the summit of Pic la Selle between
Shrubs or trees to 10–15 m with monopodial, erect 2300–2550 m on limestone, presumably above the
trunk to 40 cm diam., dioecious. Bark on tree trunks natural forest limit.
thin, exfoliating in strips or thin plates, becoming
grey-brown. Branches spreading horizontally or as-
Uses
cending, of higher order spreading or drooping in
trees, forming dense irregular or rounded crowns. No current uses have been recorded for this species;
Foliage branchlets numerous, irregularly disposed its durable wood was in demand locally in the past.
at angles between 25–45 degrees, mostly shorter
than 20 mm, slender or thicker in one variety, 0.9– 3 varieties are recognized:
1.5 mm diam., more or less quadrangular in ultimate
branchlets, persistent. Leaves decussate, decurrent, Juniperus gracilior Pilg. var. gracilior. Type:
scale-like, imbricate, with free apex on ultimate Dominican Republic: Cordillera Central, Sierra
branchlets, 1.5–2 × 0.6–1 mm, rhombic to rhombic- de la Constanza, Las Cañitas, M. Fuentes &
lanceolate; margins entire, hyaline; apex obtuse, D. Goodwin 1939 (lectotype NY).
mucronate or acute-acuminate; stomata few at base
abaxially, in one concave field adaxially; abaxial
Description
gland more or less conspicuous in basal to central
part of leaf, rounded to oval; leaf colour green. Pol- Trees. Ultimate branchlets drooping or pendulous,
len cones terminal on ultimate branchlets, subglo- spreading at 25–30 degrees, slender, 0.9–1.1 mm
bose, very small ca. 2 mm; microsporophylls 6–8, diam., more or less quadrangular. Leaves distinctly
decussate, peltate, with rounded and hyaline upper longer than wide, obtuse or acute-acuminate, gland
margin, abaxially bearing 2–3(–4) globose pollen central.
sacs. Seed cones terminal on straight, short ultimate
branchlets, maturing in one season, becoming glo-
Distribution
bose (with a single seed) or broad pyriform to near-
ly reniform, 4–5(–6) mm wide, usually soft pulpy, Hispaniola: Dominican Republic (La Vega Prov.,
resinous, reddish brown with glaucous blue bloom. near and 14 km W of Constanza; Azua Prov., Valle
Bract-scale complexes 4, decussate, the lower pair del Yaque).
small and sterile, entirely fused; sutures or bract tips TDWG codes: 81 DOM
 Juniperus gracilior Pilg. var. urbaniana (Pilg. &
Conservation
Ekman) R. P. Adams, Phytologia 78: 144. 1995.
Encroachment of human habitation and the con- Juniperus urbaniana Pilg. & Ekman, Ark. Bot. 20-A
comitant loss of habitat have reduced the three sub- (15): 9. 1926; Juniperus barbadensis L. var. urbaniana
populations known of this species considerably. Even (Pilg. & Ekman) Silba, Phytologia 56: 341. 1984;
at its greatest historically known extent it would not Juniperus barbadensis L. subsp. urbaniana (Pilg. &
have occupied more than a few km2 (AOO less than Ekman) Borhidi, Acta Bot. Acad. Sci. Hungarica
10 km2) and its population is severely fragmented. It 37 (1–4): 90. 1992. Type: Haiti: Ouest, Massif
is now known from only three localities, in each of de la Selle, Morne de la Selle, near the summit,
which fewer individuals survive. E. L. Ekman 3157 (holotype S).
420 IUCN: EN (B1+2c)
Description
Juniperus gracilior Pilg. var. ekmanii (Florin)
R. P. Adams, Phytologia 78: 144. 1995. Juniperus Decumbent or low shrubs. Ultimate branchlets
ekmanii Florin, Ark. Bot. 25-A (5): 14. 1934. Type: spreading, curved, rigid, thick, 1.3–1.5 mm diam., dis-
Haiti: Ouest, Massif de la Selle, Badaud, [“Badeau”], tinctly quadrangular. Leaves nearly as wide as long,
E. L. Ekman 3140 (holotype S). mucronate; glands often inconspicuous, near base.

Description Distribution
Trees. Ultimate branchlets spreading at 30–45 de- Hispaniola: Haiti, Massif de la Selle (Pic la Selle), near
grees, slender, 0.9–1.0 mm diam., more or less quad- the summit on limestone formation at 2300–2500 m.
rangular. Leaves slightly longer than broad; apex TDWG codes: 81 HAI-HA
acuminate or mucronate; gland conspicuous, near
base.
Conservation
Restricted to a single, small locality, from a small
Distribution
range that has probably been severely reduced since
Hispaniola: Haiti, Massif de la Selle (Morne la Selle, this variety was first discovered.
Morne la Visite?). IUCN: EN (B1)
TDWG codes: 81 HAI-HA

Juniperus horizontalis Moench, Methodus: 699.

Conservation
1794. Type: Canada: Nova Scotia, Halifax, [“prope
This variety is now limited to one or perhaps two Halifax”], M. Hultgren s.n. (neotype BM).
localities.
IUCN: CR (D) Juniperus horizontalis Moench subsp. hamptonensis
Silba, J. Int. Conifer Preserv. Soc. 11 (1): 28 (–29, 32).
2004.
Juniperus horizontalis Moench subsp. neopangaea
Silba, J. Int. Conifer Preserv. Soc. 11 (1): 28 (31). 2004.

Etymology
The species epithet refers to the growth habit close
to the ground.

Vernacular names
Creeping juniper, Waukegan juniper, Ameri-
can savin, Shrubby red cedar; Savignier, Sevigné
(French)
 normal prostrate or decumbent forms in growing
Description
more upright (to 2 m) or with an “oval” or “bonsai”
Prostrate or decumbent shrubs to 1 m, usually lower appearance. Some of the latter plants had very small
and rarely taller, spreading with rooting branches (3 mm) seed cones. The typical form was also found
over soil and rock, dioecious or rarely monoecious; in the area but said to be uncommon. Other species
stem to ca. 5 cm diam. Bark on thicker branches in the area are J. communis and J. virginiana. Such
and stems in thin flakes or wide strips, weathering growth forms, while unusual, are unlikely to be true
grey-brown. Branches creeping, spreading or whip taxa and, at the periphery of the southeastern range
shoots assurgent, higher order branches ascending of J. horizontalis, could have involved introgression
or spreading against the ground or rocks. Foliage from J. virginiana (see taxonomic notes under the
branches 3–15 cm long, often arranged upright along latter species). 421
a prostrate, rooting branch, or prostrate also, ulti-
mate branchlets densely crowded, curved, twisted
Distribution
or nearly straight, of irregular length, more or less
quadrangular in cross-section, with appressed scale Boreal and Subarctic North America, from Alaska
leaves 1–1.5 mm wide. Leaves decussate, sometimes (disjunct) to Newfoundland, more scattered further
in parts in irregular whorls of 3, imbricate, small south in the USA.
scale leaves appressed, scale leaves on ultimate TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
branchlets rhombic, 1.5–2.2 × 1–1.2 mm; margins MAN SAS 72 LAB NSC NFL-NE ONT QUE 73 COL
entire; apex acute or acuminate, transitional leaves MNT WYO 74 ILL IOW MIN NDA SDA WIS 75 MAS
common, lanceolate to acicular, distal part or only MIC NWY VER
the acuminate to pungent apex free, 3–6 mm long,
leaves on whip shoots long decurrent, to 10–13 × 2
Ecology
mm; apex caudate-pungent; abaxial stomata in 1–2
bands near base, adaxial stomata scattered from This species occurs in a variety of habitats, usually
base to apex; glands conspicuous but inactive, on on more or less open ground; on sandy beaches and
small scale leaves elliptic, on longer leaves linear, in sand dunes, on dry rocky slopes and outcrops,
leaf colour green, turning reddish purple in win- limestone ridges, in dry ‘barrens’, in grassland (prai-
ter. Pollen cones numerous, terminal, solitary, sub- ries), open bogs or ‘muskeg’ (Picea mariana / Pinus
globose to ovoid, 1.5–3 × 1–2 mm, yellowish green banksiana bog-woodland), or heathland (e.g. Em-
turning yellowish brown; microsporophylls (10–)12, petrum nigrum), or on stream banks, often forming
decussate, imbricate, peltate, with hyaline, entire or wide patches. The altitudinal range is from 10 m to
slightly erose margins and obtuse apex, bearing 3–5 1160 m a.s.l.
abaxial pollen sacs. Seed cones terminal on short,
usually curved ultimate branchlets, maturing in two
Conservation
years to irregular, then (sub)globose cones (3–)5–8
mm diam., pinkish turning brownish blue or black- IUCN: LC
ish blue. Bract-scale complexes 6, decussate, rarely
two whorls of 3, completely fused at maturity but re-
Uses
taining partly visible scale and bract margins, outer
surface smooth, usually pruinose; cone tissue soft Creeping juniper is a desirable species for horti­
pulpy and resinous. Seeds (2–)3–4 per cone, more culture in cool temperate to cold regions, since 1830
or less ovoid, often slightly curved, 3–4(–4.5) × 2–3 known in cultivation in England. There are now nu-
mm, with shallow grooves, resinous; apex obtuse, merous garden forms (cultivars) known, selected
brown, with a light hilum a third of seed length. both in Europe and North America and widely used
in gardens and parks. It makes an excellent ground
cover on sandy soils and in rockeries. Many forms
Taxonomic notes
that originated in cultivation have in the past been
John Silba (op. cit.) described two new subspecies given botanical latinized names but since they are
from a small area on Long Island, NY which are not taxa but cultivars these names should be changed
mainly based on habit that appears to differ from the to cultivar names written in inverted commas (or
 preceded by ‘cv.’), capitalized and in roman type; e.g. the seed; bract tip subapical, small, ca. 0.5 mm; sur-
Juniperus horizontalis var. viridis Grootendorst be- face smooth; scale tissue succulent, resinous. Seeds
comes J. horizontalis ‘Viridis’. All such names were single, ovoid, laterally compressed or more or less
excluded from my Monograph of Cupressaceae and flattened, 5–6 × 4 mm, shallowly grooved, pale yel-
Sciadopitys (Farjon, 2005a) which only includes taxa lowish brown.
(ranked entities classifying variation that occurs in
nature).
Distribution
Himalaya, E into high mountains of China. Bhutan;
Juniperus indica Bertol., Misc. Bot. 23: 16, t. 1. 1862. China: W Sichuan, S Xizang [Tibet], NW Yunnan;
422 India: Himachal Pradesh, Uttar Pradesh, Kashmir;
Nepal; N Pakistan; Sikkim.
Etymology
TDWG codes: 36 CHC-SC CHC-YN CHT 40 EHM-
The species epithet denotes the country of origin, i.e. BH EHM-DJ EHM-SI NEP PAK WHM-HP WHM-JK
India, from where it was first described. WHM-UT

Vernacular names Ecology

Black juniper, Wallich’s juniper; dian zang fang zhi From upper montane coniferous forest and wood-
bai (Chinese) land in pure stands, or with e.g. Abies, Pinus, Cupres-
sus torulosa, or in Betula utilis subalpine woodland,
to alpine heath and grassland and into the bare
Description
moraines and scree of the niveous zone. The altitu-
(Decumbent) shrubs, rarely small trees to 15(–20) m dinal range is from 3600 m to 4800 m a.s.l. As an
tall, dioecious; multistemmed or monopodial; trunk understorey shrub or tree in coniferous forest it is
up to 1.2 m d.b.h. Bark on larger stems exfoliating in often accompanied by J. squamata, Rhododendron
sheets or longitudinal strips, weathering dull brown. spp., Rosa, and Cotoneaster. Above the tree line it
Branches ascending or spreading, foliage branches can form juniper-rhododendron thickets, grow in
dense, short, stiff and spreading or erect, forming Kobresia-Stipa turf with dwarfed alpine shrubs (e.g.
a dense, broad pyramidal to eventually rounded Rhododendron, Salix, Juniperus squamata), or occur
or irregular crown. Foliage branchlets ultimately scattered on moraines and consolidated scree slopes
stout, quadrangular, sometimes more or less terete, of granite or gneiss or other metamorphic acidic
1.2–1.5 mm wide, covered with appressed or some- rock, at the highest altitudes exclusively on S-facing
times spreading leaves, persistent. Leaves on mature slopes. The climate is high montane to alpine with a
plants eventually scale-like, decussate, sometimes pronounced monsoon phase delivering heavy pre-
3-whorled, imbricate, decurrent, 1.2–2 × 1–1.2 mm, cipitation (much as snow) from May to October.
triangular-rhombic, obtuse; margins entire; scale
leaves amphistomatic, stomata in 2 inconspicuous
Uses
lines on each side mostly near base; glands central
in a groove, elliptic or oblong, sometimes absent; Black juniper is probably rare in cultivation under
leaf colour green or slightly glaucous green. Pollen either J. wallichiana or J. pseudosabina; the first being
cones numerous, solitary, terminal on short branch- a synonym of J. indica and the latter a closely related
lets, subglobose to ovoid-globose, 2–3 mm long; species from Central Asia. The recently discovered
microsporophylls 6–8, decussate, peltate-cordate, variety J. indica var. caespitosa is a decumbent form
with rounded entire hyaline margins, bearing 2–3 suitable as ground cover in rock gardens, but not yet
abaxial pollen sacs. Seed cones terminal on short in cultivation. The growth habit of J. indica var. indi-
erect branchlets, maturing in the second season to ca is erect, but varies from shrub to small tree. Both
subglobose or ovoid, (4.5–)5–13 × 4–8 mm, lustrous forms are commendable for horticulture, but slow
blue-black or brownish black, soft cones. Bract- growing. In the Himalayas, where Black juniper is
scale complexes 3–6, decussate or (2×) 3-whorled, common and widespread, its wood is used for fuel
entirely fused, sometimes incompletely covering
and branches and foliage are burned as incense in the tall, erect form of the species and within the up-
Buddhist temples. permost belt of subalpine forest dominated by Abies
and Larix.
2 varieties are recognized:
Conservation
Juniperus indica Bertol. var. indica. Type: Illustra-
tion in Bertoloni, Misc. Bot. 23: 16, t. 1. 1862 (lecto- IUCN: LC
type).

Juniperus wallichiana Hook. f. & Thomson ex Juniperus jaliscana Martínez, Anales Inst. Biol.
E. Brandis, Forest Fl. NW & Central India: 537. 1874. Univ. Nac. México 17 (1): 69. 1946. Type: Mexico: 423
Jalisco, Talpa de Allende, Cumbre Blanca,
M. Martínez & A. Q. Gonzalez 7002 (holotype
Description
MEXU).
Erect shrubs to small trees to 15(–20) m tall. Foli-
age branches spreading or more or less erect, dense
Etymology
with short branchlets. Seed cones when mature
mostly (broadly) ovoid, 8–13 × 5–8 mm, blue-black The species epithet refers to the State of Jalisco in
or brownish black. Mexico.

Distribution Vernacular names

As the species. Cedro (Spanish)

Conservation Description
IUCN: LC Small trees to 10 m tall, dioecious or monoecious;
trunk erect, more or less straight. Bark on trunk fi-
Juniperus indica Bertol. var. caespitosa Farjon, brous, soft, exfoliating in long thin strips, orange-
Monogr. Cupressaceae & Sciadopitys: 313. 2005. brown to reddish brown, weathering grey. Branches
Type: Nepal: Dhaulagiri Himal, Dolpo, Mugu ascending or spreading, in younger trees nearly erect,
Karnali, S of Mugu, S. Miehe 99-03001 (holotype K). forming a conical crown, later a rounded crown. Fo-
liage branches numerous, dense, slender and lax,
ultimate branchlets more or less curved, of unequal
Description
length up to 15 mm, thin, 1.1–1.3 mm wide, more or
Decumbent or ascending shrubs 50–100 cm tall. Fo- less quadrangular in cross-section. Leaves all scale-
liage branches (nearly) erect, very dense with short like, decussate, not or slightly imbricate, persistent
branchlets. Seed cones when mature (sub)globose on 3–4 orders of branching, on ultimate branchlets
to broadly ovoid, (4.5–)5–8 × 4–6.5 mm, blue-black. ovoid-oblong, 1–1.5 × 0.7–0.8 mm (growing to 4–5
mm long and 2–2.5 mm wide with a caudate, acute
apex on older lower order branchlets), slightly in-
Distribution
curved; margins minutely or remotely denticulate to
Nepal, S Xizang [Tibet], Bhutan; probably elsewhere nearly entire; apex free, acute-acuminate; amphisto-
and as yet incompletely known. matic, nearly all stomata adaxially; glands central,
TDWG codes: 36 CHT 40 EHM-BH NEP linear or oblong, inactive; leaf colour grey-green.
Pollen cones terminal, not seen. Seed cones termi-
nal, solitary on short, curved branchlets, maturing
Ecology
to subglobose, irregular cones of 5–7.5 mm diam.,
This is a decumbent form that is generally associ- brown, variously glaucous. Bract-scale complexes 6,
ated with tree line habitat and above, but in some decussate, with protruding, recurved bract tips up to
localities it has been found growing amongst both 1 mm long, pulpy tissue hardening when drying and
 shrinking. Seeds 5–8(–10) per cone, of unequal size,
Vernacular names
the largest 4 × 2.5 mm, narrowly ovoid, more or less
acute on both ends, light brown, with yellow resin ta zhi yuan bai (Chinese)
drops at chalazal end, micropylar protusion some-
times persistent.
Description
Trees to 20 m, monoecious; trunk up to 1.2 m d.b.h.
Distribution
Bark on trunk stringy, exfoliating in longitudi-
Mexico: S Durango, NW Jalisco, only known from nal strips, light brown or greyish brown. Branches
two localities. spreading from trunk, foliage branches drooping
424 TDWG codes: 79 MXE-DU MXS-JA to pendulous, branchlet systems often tapering
with distal branchlets successively shorter. Foli-
age branchlets straight or slightly curved, ultimate
Ecology
branchlets terete or quadrangular, stout, 1.2–1.5 mm
The ecology of this species is not well known. Some diam., covered with green scale leaves, persistent.
specimens have been collected in goat pasture, with Leaves mostly decussate, triangular-ovate, 1.5–3.5 ×
the grasses frequently burned, on steep, weathered 1–1.5 mm; margins entire; apex acute or sometimes
granitic slopes, i.e. disturbed secondary vegetation. obtuse; stomata restricted to concave distal part of
The altitudinal range is reported to be from 430 m to scale leaves; abaxial gland often inconspicuous, in
2670(?) m. The divergent altitudes of the two locali- lower part of leaf or near the base, orbicular to ob-
ties are remarkable; whereby it must be stated that long, convex; leaf colour green. Pollen cones soli-
the high altitude was reported for the locality that tary and terminal on ultimate branchlets, globose
has not been identified. to ovoid, 2–3 mm long; microsporophylls (6–)8–10,
decussate, peltate, with rounded margin, bearing
2–3 abaxial pollen sacs near lower margin. Seed
Conservation
cones terminal, initially (sub)erect on short branch-
This species is only known from two localities, one lets, becoming pendulous with growing branchlets,
of which, mentioned by Martínez (op. cit.) as the maturing in two seasons, turning from (brownish)
place in Durango where C. E. Blanco collected it, green to bluish or purplish black when ripe, ovoid
has not been found back recently, nor traced on the (in green cones with acute apex), 8–10(–12) × 6–8(–
map. Juniperus jaliscana occurs in grazed secondary 9) mm, smooth, lustrous in ripe, blackish cones or
vegetation as well as in woods (at the higher altitude slightly glaucous. Bract-scale complexes 6, decus-
in Durango). It is listed as Endangered even though sate; sutures visible at least until maturity; bract tips
it is likely that a thorough exploration of the relevant small, acute, protruding near apex. Scale tissue soft
regions would turn out more individuals or even pulpy, resinous. One seed per cone, ovoid or ellip-
populations. soid with acute base and apex, more or less with two
IUCN: EN (B1+2c) opposite ridges, irregularly grooved, 7–10 mm long,
light brown, with darker resin pits near base and 1–2
pits near middle.
Juniperus komarovii Florin, Acta Horti Gothob.
3: 3. 1927. Type: China: Sichuan, N Sichuan,
Distribution
Sangarmai, [“Sankar-vou-mâ”], H. Smith 4402
(holotype GB). China: S Gansu, SE Qinghai, NW Sichuan.
TDWG codes: 36 CHC-SC CHN-GS CHQ
Etymology
Ecology
The species epithet commemorates the Russian Bot-
anist Vladimir L. Komarov (1869–1945), compiler of This species has been found in deep gorges and
the multi-volume Flora of the USSR. on rocky or grass-covered slopes in the semi-arid
mountains and plateaus of S Gansu, SE Qinghai and weakly quadrangular in cross section, covered with
NW Sichuan, where it is associated with few other scale leaves, persistent. Leaves in alternate whorls of
trees and mainly restricted to S-facing slopes. The 3, less often decussate on ultimate branchlets, im-
altitudinal range is from 2110 m to 4000 m a.s.l. N- bricate, scale-like, 1–2(–3) × 0.6–1.5 mm on lateral
facing slopes are covered with spruce forest (Picea branchlets, decurrent, rhombic to broadly rhombic;
asperata). apex appressed or sometimes free, acute/acuminate;
margins minutely denticulate; stomata on abax-
ial side limited to decurrent leaf base, on adaxial
Conservation
surface in two tapering bands; glands more or less
IUCN: LC conspicuous, large, elliptical, slightly depressed,
exudate often present; leaf colour yellowish green 425
or light green. Pollen cones numerous, terminal,
Uses
solitary, ellipsoid or ovoid-oblong, 3–4 × 2–2.5 mm;
No uses have been recorded for this species, but its microsporophylls 8–10, decussate, peltate, acute,
wood and foliage may be used locally for firewood with minutely denticulate upper margins and 3–4
and incense burning. In cultivation it is only known abaxial pollen sacs. Seed cones terminal on straight,
in a few arboreta and these are recent introductions. very short branchlets, maturing to purplish orange
or reddish brown with glaucous or blue bloom in
one season, subglobose to ovoid-globose, 5–8 mm
Juniperus monosperma (Engelm.) Sarg., Silva diam., internally soft pulpy, succulent, yellowish to
N. Amer. 10: 89. 1896. Juniperus occidentalis Hook. orange. Bract-scale complexes 4–6, entirely fused,
var. monosperma Engelm., Trans. St. Louis Acad. decussate, bract apices prominently exserted, sur-
Sci. 3: 590. 1878. Type: USA: Colorado, Fremont face smooth, rugose when dry, glaucous. Seeds 1(–2)
Co., Canon City, G. Engelmann s.n. (lectotype MO). per cone, broadly ovoid, 4–6 × 3–4 mm, base trun-
Pl. 17 cate; apex acutish, shallowly grooved and ridged on
sides, not resinous, light yellowish brown to chest-
nut brown, with a large, lighter lobed hilum.
Etymology
The species epithet means ‘with a single seed’, but
Distribution
this species is not strictly monospermous, as are
some others. USA: Arizona, S Colorado, S Nevada, New Mexico,
W Oklahoma, NW Texas; Mexico: Chihuahua.
TDWG codes: 73 COL 74 OKL 76 ARI NEV 77 NWM
Vernacular names
TEX 79 MXE-CU
One-seed juniper, Cherrystone juniper, West Texas
juniper; Sabina (Spanish)
Ecology
In Pinyon-Juniper woodland, with Pinus edulis, Ju-
Description
niperus osteosperma, J. scopulorum, and in grassland
Large shrubs to 6 m, dioecious, rarely monoecious; with Opuntia spp., Yucca spp. on dry, rocky soils of
multistemmed or rarely with a short trunk and a intermountain basins, or on rock outcrops, buttes
maximal diam. of 30 cm. Bark of thick branches and mesas of limestone, sandstone or igneous rock.
and trunk fibrous, exfoliating in strips, weathering The altitudinal range is from 970 m to 2200 m a.s.l.
ash-grey. Branches numerous, ascending from base The climate is continental semi-arid, with summer
or nearly erect, often curved, those of higher orders rainfall mainly arriving with erratic and local storms
ascending or spreading, forming a broad, rounded from the Gulf of Mexico.
or more irregular and open crown. Foliage branches
numerous, irregularly disposed but not pendulous,
Conservation
ultimate branchlets spreading to erect, slender and
stiff, up to 15 mm long, 1.2–1.9 mm thick, subterete or IUCN: LC
 8

4
426

6
7

5 3 2

Pl ate 17. Juniperus monosperma. 1. Habit of shrub. 2. Branch with foliage. 3. Branchlet with leaves.
4. Leaves with gland. 5. Pollen cone. 6, 7. Seed cones, one opened to show seed. 8. Seeds.
 stomata scattered adaxially, covered by thick cuticu-
Uses
lar wax; glands central, elliptic or nearly linear, or
The wood is useful as firewood and for fence posts. absent on many smallest leaves; leaf colour light to
The fibrous bark was woven into mats and cloth by dark green or glaucous green. Pollen cones termi-
Native Americans (Apache, Comanche, Ute). One- nal, solitary, oblong, 3–4 × 2–2.5 mm, more or less
seed juniper is of no particular interest to horticul- quadrangular in cross section; microsporophylls
turists probably because of its growth habit, often 10–12, decussate, peltate, often keeled, bearing 3–4
not ‘knowing’ whether it wants to be a shrub or a abaxial pollen sacs. Seed cones terminal on very
tree. short branchlets; mature cones globose or subglo-
bose, 6–9(–10) mm diam., lustrous, purplish black
or purplish red, sometimes dull orange-brown, usu- 427
Juniperus monticola Martínez, Anales Inst. Biol. ally with a glaucous bloom. Bract-scale complexes
Univ. Nac. México 17 (1): 79. 1946. Type: Mexico: 6, completely fused, forming a soft pulpy, succulent,
Hidalgo, Mineral del Monte, Rancho del Guajolote, resinous cone with smooth surface and minutely
C. A. Ehrenberg s.n. (isotype MO). exserted (lower) bract tips. Seeds (2–)3–7(–9) per
cone, angular ovoid, irregular but more or less flat-
Juniperus monticola Martínez var. monticola f. com- tened, 3–5 × 2–3 mm, small relative to cone size,
pacta Martínez, Bol. Soc. Bot. México 7: 19. 1948; with a small light hilum at or near base, grooved or
Juniperus compacta (Martínez) R. P. Adams, Phyto- pitted proximally, with resin in pits, lustrous (dark)
logia 89 (3): 368. 2007. chestnut-brown.

Etymology Taxonomic notes

The Latin species epithet means ‘mountain-dwell- Recently, Adams (op. cit.) raised Juniperus monti-
ing’. cola var. monticola f. compacta to the rank of spe-
cies: J. compacta (Martínez) R. P. Adams, based on
analysis of DNA sequence data and biochemis-
Vernacular names
try. No morphological characters seem to separate
Mexican juniper; Sabina, Savino (Spanish); Tláscal J. monticola from J. compacta other than the habit
(Aztec) mentioned by Martínez in his original description.
Whilst it may be so that these biochemical differ-
ences exist (and even could turn out to be consistent
Description
with wider sampling), it remains to be demonstrated
Decumbent shrubs to small trees 6–10 m, dioecious; that it is taxonomically informative to base species
trunk prostrate and tortuous to erect, usually forked concepts on such data, creating, as Adams admits,
near base or branching low, to 60 cm diam. Bark on “cryptic species”.
larger trunks becoming fibrous, to ca. 1 cm thick,
exfoliating in long strips. Branches spreading or as-
Distribution
cending to nearly erect; forming a variable crown
from decumbent to broadly rounded and dense. Mexico: Guerrero, Hidalgo, Jalisco, México, México
Foliage branches often in tapering tufts, ultimate D. F., Michoacan, Nuevo León, Puebla, Veracruz;
branchlets numerous, straight or curved, (3–)5–20(– Guatemala: Huehuetenango.
22) mm long, quadrangular in cross section, often TDWG codes: 79 MXC-DF MXC-ME MXC-PU MXE-
thick on plants from above ca. 3000 m (1–2-1.7 mm), NL MXE-SL MXG-VC MXS-JA MXS-MI MXS-GR 80
other localities 1.2–1.4 mm diam., persistent. Leaves GUA
decussate, imbricate, closely appressed, appearing in
four rows, or in some plants more irregularly ori-
Ecology
ented and less closely appessed, on ultimate branch-
lets broadly rhombic, 0.8–1.4 × 1–1.2 mm, distinctly The large shrub to small tree form of this species is
keeled; margins minutely hyaline-erose to serrulate; occasionally found in Pinyon-Juniper woodland, or
apex obtuse; few stomata abaxially on lower margin, in other open pine forest; it has also been reported
 from “Ficus religiosa forest”, with an understorey of 5 cm long, (1.2–)1.5–2 mm thick, covered with scale
Ribes, Rosa and Rubus (herbarium collection Zanoni leaves, persistent. Leaves in alternate whorls of 3, less
2718, Mexico, Federal District, 2895 m). The decum- often decussate on ultimate branchlets, closely ap-
bent shrub form is common at or above the tree line pressed, scale-like, 2–3 × 2 mm on ultimate branch-
(usually with Pinus hartwegii) on ridges of basalt or lets, ovoid-rhombic; margins minutely denticulate,
consolidated scree, with grasses e.g. Calamagrostis acute or obtuse; stomata abaxially near base and in
and Festuca, shrubs such as Ribes, and some alpine two tapering bands adaxially; glands conspicuous,
herbs, or on rocks and cliffs to the limit of vegetation central, oval, often active with white-drying resin-
on Mexico’s highest volcanoes. The altitudinal range ous exudate; leaf colour dark green. Pollen cones
of the species is 2000–4270 m a.s.l. Despite high numerous, terminal on short ultimate branchlets,
428 precipitation on these mountains, the environment subglobose to ovoid-oblong, 3–5 × 2–3 mm; mi-
is decidedly xeric due to the porosity of volcanic crosporophylls (10–)12–16(–18), mostly decussate,
substrates, but snowmelt may provide a relatively peltate, bearing 3 abaxial pollen sacs near lower
reliable, slow-release moisture source in spring and margin. Seed cones terminal on ultimate branch-
early summer. lets, growing in two seasons from purplish red to
blue or purple with a glaucous bloom, subglobose
to ellipsoid, (6–)7–9(–10) × (5–)6–8 mm, fleshy or
Conservation
pulpy, more or less resinous, eventually dry. Bract-
IUCN: LC scale complexes (4–)6, decussate or in two alternate
whorls of 3, entirely fused, smooth, rugose when
dry, with small triangular bract apices protruding.
Uses
Seeds 1–2 per cone, ovoid or semi-ovoid, 5–6(–7)
This species is not in cultivation, but the decumbent × 4–5 mm, with shallow grooves and resinous pits,
alpine form would probably be suitable for rock gar- yellowish brown.
dens and withstand low winter temperatures.
Distribution
Juniperus occidentalis Hook., Fl. Bor. Amer. 2 (10): W USA: California, W Idaho, Nevada, Oregon,
166. 1838. Washington.
TDWG codes: 73 IDA ORE WAS 76 CAL NEV
Etymology
Ecology
The Latin species eithet means ‘from the west’.
In the northern part of its range (var. occidenta-
lis) Western juniper is often forming single species
Vernacular names
stands, or it is associated with Pinus ponderosa. Here
Western juniper, Sierra juniper (var. australis) it is most frequently found on flood plains, terraces,
and grassy plateaux and uplands. In E Oregon it is
also spreading onto abandoned agricultural fields
Description
and rangeland, partly as a result of a decreasing
Trees 10–15(–20) m tall, to 1.5–2.5 m diam., mono­ frequency of wildfires. In the Sierra Nevada (var.
ecious or dioecious; trunk monopodial or with sev- australis) it can grow with Pinus jeffreyi, P. albi­
eral massive stems low above the ground. Bark on caulis, P. monticola, P. contorta, Abies magnifica,
trunk fissured, fibrous, exfoliating in long strips, Calocedrus decurrens, and Tsuga mertensiana. The
orange-brown to (reddish) brown. Branches thick, most common understorey shrub in its entire range
ascending or spreading, often curved or contorted, is Seriphidium tridentatum (Artemisia tridentata);
the foliage branches forming dense rounded tufts at in the north also Chrysothamnus spp., Purshia tri-
ends of heavy main branches, forming a pyramidal dentata, Ribes cereum, in the south Arctostaphylos
crown in young trees, becoming rounded and irreg- spp., Ceanothus sp., and several others. The altitu-
ular in old trees. Foliage branchlets numerous, stout, dinal range is 200–1200 m a.s.l. (var. occidentalis)
subterete to quadrangular, ultimate branchlets to and 1980–3100 m a.s.l. (var. australis). In coniferous
forest it usually occurs where rock outcrops cause date with accuracy, are on protected public lands,
shallow soils, at higher altitudes in the Sierras it can where cutting trees and grazing livestock are either
usually be found among granite boulders or even in prohibited or strictly regulated. They are of great
crevices on bare granite domes. In the north it grows aesthetic and scientific value.
on xeric soils, often derived from volcanic rock, or IUCN: LC
from non-calcareous sediment. The climate ranges
from semi-arid in the rain shadow of the Cascades Juniperus occidentalis Hook. var. australis (Vasek)
(15–45 cm p/a) in the north to mesic in the Sierras, P. Lebreton & N. H. Holmgren, Intermountain Fl.
with precipitation mostly in the form of winter snow. 1: 239. 1972. Juniperus occidentalis Hook. subsp.
australis Vasek, Brittonia 18: 352. 1966; Juniperus
grandis R. P. Adams, Phytologia 88 (3): 306. 2006. 429
Uses
Type: USA: California, San Bernardino Co., San
The wood of this juniper is still used locally for Bernardino Mts., along Polique Canyon Road to
firewood, fence poles, corrals, etc. Its very durable Holcomb Valley, F. C. Vasek 610929-38 (holotype
wood is also used for special purposes such as furni- RSA). Fig. 140
ture, interior panelling and decorative applications.
Horticultural use is rare, but the species has value
Description
for landscaping of disturbed sites. There are a few
cultivars known, one of these has conspicuously sil- Trees mostly dioecious, occasionally monoecious;
very glaucous foliage and is called ‘Sierra Silver’ and, bark orange-brown or reddish brown. Scale leaves
at least for a time, retains the columnar or pyramidal mostly in whorls of 3. Cotyledons 2–4 in embryos,
habit of younger plants. 2–4 in seedlings.

2 varieties are recognized:

Distribution
Juniperus occidentalis Hook. var. occidentalis. Type: W USA: California, extreme W Nevada.
USA: Washington, Columbia River, [“Common on TDWG codes: 76 CAL NEV
the higher parts of the Columbia, at the base of the
Rocky Mountains”], D. Douglas s.n. (holotype K).
Conservation
IUCN: LC
Description
Trees mostly monoecious (“submonoecious”); bark
“brownish”. Scale leaves decussate or in whorls of 3. Juniperus osteosperma (Torr.) Little, Leafl. W. Bot.
Cotyledons 2–4 in embryos, usually 2 in seedlings. 5: 125. 1948. Juniperus tetragona Schltdl. var. osteo­
sperma Torr., Pacif. Railr. Rep. 4 (5) [no. 4]: 141.
1857; Juniperus californica Carrière subsp. osteo­
Distribution
sperma (Torr.) E. Murray, Kalmia 12: 21. 1982. Type:
W USA: N California, W Idaho, NW Nevada, Or- USA: Arizona, Coconino Co., Bill Williams Mt.,
egon, S Washington. J. M. Bigelow s.n. (lectotype NY).
TDWG codes: 73 IDA ORE WAS 76 CAL NEV
Etymology
Conservation
The species epithet means ‘with bony seed’, presum-
Of conservation concern, mostly for other reasons ably referring to the hardness of the seeds.
than species survival in the strict sense, are the very
ancient specimen trees of both recognized varieties
Vernacular names
which occur in the Cascades and especially in the
Sierra Nevada (var. australis). Many of these trees, Utah juniper, Bigberry juniper; Sabina morena
possibly several thousand years old but difficult to (Spanish)
 TDWG codes: 73 COL IDA MNT WYO 76 ARI CAL
Description
NEV UTA 77 NWM
Arborescent shrubs or small trees 3–6(–12) m tall,
monoecious or rarely dioecious; trunk to 1 m diam.
Ecology
near base, with several stems low above ground. Bark
on trunk scaly, fibrous, exfoliating in long strips, This species is a codominant in the widespread
(reddish) brown weathering grey. Branches thick, Pinyon-Juniper woodland of the Intermountain
spreading and ascending, often curved or contorted, Region of the American West, with Pinus edulis,
the foliage branches forming tufts at ends of main P. monophylla (in the SW of its range), P. cembroides
branches; crown becoming rounded and irregular in (in the S of its range), Juniperus scopulorum, J. oc-
430 old trees. Foliage branchlets numerous, thick, sub- cidentalis (in the W of its range), Seriphidium tri­
terete or sometimes quadrangular, ultimate branch- dentatum (Artemisia tridentata), Chrysothamnus
lets to 3 cm long, 2–2.5 mm thick, covered with spp., Quercus gambelii, and Ephedra viridis, or in
scale leaves, persistent. Leaves in alternate whorls pure stands at the lower altitudinal limits of the
of 3, sometimes decussate on ultimate branchlets, community. The altitudinal range is 460–2700 m
closely appressed, scale-like, 1.5–3.5 × 1–2.5 mm a.s.l. Juniperus osteosperma is abundant on rocky or
on ultimate branchlets, ovoid-rhombic to rhom- gravelly alluvial fans and hillsides, as well as on bare
bic; margins minutely denticulate, acute or obtuse; sandstone or shale where it finds crevices in the rock
abaxially a few stomata near base and in two taper- to grow in. It is one of the most drought resistant
ing bands adaxially; glands inconspicuous, central, junipers in North America.
usually inactive; leaf colour green with whitish wax.
Pollen cones numerous, terminal on short ultimate
Conservation
branchlets, subglobose to ovoid, 3–4.5 × 2–2.5 mm;
microsporophylls (10–)12–16, mostly decussate, IUCN: LC
peltate, bearing 3–4 abaxial pollen sacs near lower
margin. Seed cones terminal on short branchlets,
Uses
growing in two seasons from blue-green or purplish
to light brown with a glaucous white bloom, subglo- In many arid places on the Colorado Plateau and
bose to ovoid-ellipsoid, 7–13(–15) × 6–12 mm, pulpy elsewhere in its extensive range, this species of juni-
or fibrous, more or less resinous, maturing dry and per is the most important small tree providing im-
hard. Bract-scale complexes 4–6, usually decussate portant habitat for wildlife. Its use for firewood is
or sometimes (in part) in whorls of 3, entirely fused, now incidental and its contorted habit when grow-
smooth, slightly rugose when dry, with triangular ing older precludes most other uses of its wood. In
bract apices protruding from crescent ridges. Seeds regions with dry and hot summers it could be used
1–2 per cone, subglobose or broadly ovoid, 4–10 × in gardens; it is very hardy as it occurs naturally in
4–9 mm, with shallow grooves and resinous pits and continental climate and at high altitudes.
a large proximal hilum, light brown.

Juniperus oxycedrus L., Sp. Pl. 2: 1038. 1753.

Taxonomic notes
Genetic research (DNA) undertaken recently (Terry
Etymology
et al., 2000) has produced evidence for introgression
of J. occidentalis into J. osteosperma; this is possibly The species epithet is a combination of the Greek
linked with the recent spread of the former species ‘oxy’ (from oxus = sharp) and cedrus, the classical
into the west of the range of J. osteosperma. (Latin) word for fragrant conifer wood.

Distribution Vernacular names

W USA: Arizona, E California, Colorado, S Idaho, S Prickly juniper; Oxycèdre, Genévrier cade, Gené­
Montana, Nevada, W New Mexico, Utah, Wyoming. vrier epineux (French); Kade, Enebro de bajas
­rochas (Spanish); Katran Ardiçi (Turkish)
 TDWG codes: 12 BAL COR FRA-FR FRA-MO POR
Description
SAR SPA-AN SPA-GI SPA-SP 13 ALB BUL GRC ITA-
Shrubs or small trees to 10(–15) m tall, dioecious; IT ITA-SM ITA-VC KRI ROM SIC-MA SIC-SI TUE
multistemmed or monopodial to 50–100 cm diam., YUG-BH YUG-CR YUG-KO YUG-MA YUG-MN YUG-
low branching, sometimes prostrate with ascend- SE YUG-SL 14 KRY 20 ALG LBY MOR-MO MOR-SP
ing stems. Bark on trunks with long, soft strips, TUN 33 NCS TCS 34 CYP EAI IRN IRQ LBS-LB LBS-SY
brown turning grey. Branches numerous, spread- PAL-IS PAL-JO TUR
ing or ascending. Foliage usually dense, with short,
spreading or sometimes longer, subpendulous,
Ecology
‘jointed’ ultimate branchlets, persistent, forming a
dense, irregular and broad, sometimes pyramidal A common shrub in Mediterranean sclerophyll 431
or conical crown. Leaves in alternating whorls of 3, scrubland (maquis, garrigue); also in dry woodland
non-decurrent, spreading 60–90 degrees at nodes with Pinus spp., Carpinus betulus, Quercus ilex and
3–5(–10) mm apart, rigid, jointed to leaf part ad- other oaks, Quercus-Lentiscus scrub, as well as in
nate with shoot, subulate to linear-acicular, some- montane and wetter forest with Cedrus libani, Pinus
times narrowly ovate (‘boat-shaped’) or narrowed nigra, Juniperus foetidissima, and J. excelsa. The al-
towards base as well as apex, (4–)10–20(–25) × 1.1–3 titudinal range is 1–2200 m a.s.l. It occurs on dry,
mm, straight; margins entire; apex from more or stony slopes in thin soils over all kinds of rock from
less acute to acute-pungent; epistomatic, stomata calcareous to siliceous and serpentine, but uncom-
in two conspicuous whitish bands separated by a monly on sand dunes; in pastures at higher alti-
midrib, bordered by green margins as wide as mid- tudes it is usually a sign of overgrazing. It is largely
rib, abaxial face (weakly) keeled or ridged, green to restricted to regions with a Mediterranean climate,
dark green. Pollen cones axillary, solitary, 2–3 per but in the Balkans and the Iberian Peninsula it may
leaf whorl, subglobose to ovoid, 3–4 × 2–3 mm, yel- occur in more continental conditions. The subspe-
lowish, turning brown; microsporophylls 8–12, in cies badia and oxycedrus are tolerant of frost, un-
alternating whorls of 3, peltate, with entire margins like the subspecies macrocarpa and transtagana.
and acuminate apex, bearing 4–6 abaxial pollen The latter two are largely restricted to coastal dunes
sacs. Seed cones axillary on short (1–2 mm) dwarf or old, vegetated beaches on sand, where they oc-
shoots with whorls of minute scale leaves, maturing cur in open pine woods (Pinus halepensis, P. brutia,
in 2 years. Mature cones globose or ovoid-globose, P. pinea) as well as in scrub (garrigue, maquis) or
8–20(–23) mm long, with 1(–2) whorls of 3 com- associated with coastal grasses.
pletely fused bract-scale complexes, only the upper
whorl fully developed, with 3 sutures conspicuous
Uses
on distal half of cone; bract tip hidden or very small;
surface smooth or rugose; tissue dry pulpy, more or Prickly juniper is suitable for cultivation as an or-
less resinous; immature cones green, hard; mature namental in southern Europe where a number of
cones orange-brown to purplish brown, sometimes cultivars, especially with more pendulous foliage,
glaucous or pruinose. Seeds 2–3 per cone, triangu- are commonly planted in gardens and parks. Essen-
lar-ovoid, often slightly flattened, 5–12 × 4–8 mm, tial oils are extracted from the branches and leaves,
brown. especially in France and Turkey. This ‘oil of cade’ is
used for medicinal puposes.
Distribution
4 subspecies and 1 variety are recognized:
Mediterranean Region, around the Black Sea and
Middle East: Albania, Algeria, Bosnia-Herzegovi- Juniperus oxycedrus L. subsp. oxycedrus. Type:
na, Bulgaria, Caucasus, Croatia, Cyprus, France, Spain: [“Habitat in Hispania, G. Narbonensi”],
Greece, Iran (Azerbajian), Irac, Israel, Italy, Jordan, Herb. Burser XXV: 67 (lectotype UPS).
Lebanon, Libya, Macedonia, Malta, Morocco, Por-
tugal, Romania, Serbia, Spain, Syria, Transkaukasus, Juniperus oxycedrus L. var. oxycedrus f. parvifolia
Tunisia, Turkey, Ukraine: Krym [Crimea]. Novák, Preslia 4: 53. 1926; Juniperus oxycedrus L. var.
 parvifolia (Novák) Jovan., in Saric (ed.), Fl. Serbia 1: p­ allasiana, growing on rocky slopes at 800–1400 m
218. 1992. a.s.l.
Juniperus oxycedrus L. var. fastigiata Jovan., in Saric
(ed.), Fl. Serbia 1: 413. 1992.
Conservation
Juniperus deltoides R. P. Adams, Phytologia 86 (2):
47. 2004. IUCN: NE

Juniperus oxycedrus L. subsp. badia (H. Gay)

Description
Debeaux, Fl. Kabylie: 412. 1894. Juniperus oxycedrus
Shrubs, or small trees to 8 m, with spreading or L. var. badia H. Gay, Assoc. Franç. Avancem. Sci.
432 a­ scending branches; foliage branchlets variable but Compt. Rend. 1889: 501. 1889. Type: Algeria: 20 km
ultimate branchlets not subpendulous. Leaves 6–25 S of Medea [“l’oxycèdre des Haouaras”], H. Gay s.n.,
× 1.1–2 mm, spreading at 60–90 degrees from shoot. 17 June 1888 (holotype G?).
Seed cones globose, 6–13 mm diam., maturing to
orange- or reddish brown, rarely more or less glau-
Description
cous.
Pyramidal shrubs or trees to 15 m tall; branch-
es spreading, ultimate branchlets subpendulous.
Distribution
Leaves linear-acicular, or ‘boat-shaped’ in shortest
As for the species. leaves, (8–)12–20 × 1.2–2 mm, acuminate or acute-
pungent (short and subobtuse leaves often occur on
male plants). Seed cones globose, 10–13 mm diam.,
Conservation
yellowish brown and more or less glaucous when
IUCN: LC growing, purplish brown at maturity.

Juniperus oxycedrus L. subsp. oxycedrus var.

Distribution
spilinanus Yaltirik, Eliçin & Terzioglu, Turkish
J. Bot. 31 (1): 38. 2007. Type: Turkey: W Anatolia, N Algeria; also reported from E Portugal and central
Manisa, Spildagi N. P., F. Yaltirik et al. 13371, 28 May Spain.
1995 (holotype KATO). TDWG codes: 12 POR SPA-SP 20 ALG

Description Conservation
Dioecious, prostrate shrubs up to 0.5–0.6 m tall. IUCN: LC
Leaves lanceolate, patent, green, with two glaucous
bands adaxially (upperside), 6–10 × 1–1.5 mm, acu- Juniperus oxycedrus L. subsp. macrocarpa (Sibth.
minate-mucronate. Seed cone when ripe globose, & Sm.) Ball, J. Linn. Soc. London, Bot. 16: 670.
to 10 mm diam., reddish brown. Seeds usually 3 per 1878. Juniperus macrocarpa Sibth. & Sm., Fl. Graeca
cone, free within the pulpy cone tissue. Prodr. 2: 263. 1816. Type: Greece: [locality not
stated], J. Sibthorp s.n. (lectotype OXF). Fig. 141
Distribution
Description
Turkey, W Anatolia (Spildagi National Park).
TDWG codes: 34 TUR Shrubs or small trees with spreading branches, foli-
age branches short, thick and rigid. Leaves spread-
ing at nearly right angles from shoot, subulate, 2–3
Ecology
mm wide at base, very rigid. Pollen cones 4 × 3
This prostrate variety has been found in forest clear- mm when closed. Seed cones (12–)15–23 mm long,
ings among Pinus brutia and Pinus nigra subsp. ovoid-globose or globose, smooth or rugose, with
scale margins often prominent, turning from orange Juniperus phoenicea L., Sp. Pl. 2: 1040. 1753.
or yellowish to brown or purplish brown to purplish
red, pruinose when ripe.
Etymology
The species epithet refers to Phoenicia, the classical
Distribution
name for the famous sea-faring nation on the North
Mediterranean and Black Sea coasts: Albania, Alge- African coast (now part of Tunisia).
ria, Bosnia-Herzegovina, Bulgaria, Croatia, France,
Greece, Italy, Malta, Morocco, Spain, Turkey,
Vernacular names
Ukraine: Krym [Crimea].
TDWG codes: 12 BAL COR FRA-FR SAR SPA-SP 13 Phoenician juniper; Genévrier rouge (French); Arâr 433
ALB BUL GRC ITA-IT KRI SIC-MA SIC-SI TUE YUG- (Morocco)
BH YUG-CR 20 ALG LBY MOR-MO MOR-SP 34 EAI
Description
Conservation
Arborescent shrubs or small trees to 8–12 m tall,
IUCN: LC mono­ecious or rarely dioecious; trunk to 1 m diam.
near base, monopodial or with several stems low
Juniperus oxycedrus L. subsp. transtagana Franco, above ground. Bark on trunk and larger branches
Fedde’s Repert. Sp. Nov. Regni Veg. 68 (3): 166. scaly, exfoliating in long strips, weathering grey.
1963. Juniperus oxycedrus L. var. transtagana Branches spreading and assurgent, often curved,
(Franco) Silba, Phytologia Mem. 7: 35. 1984. Type: the foliage branches forming tufts at ends of main
Portugal: Ribatejo, Portela de Sacavem, Alcochete, branches; crown more or less conical in young trees,
[“in arenosis trans Tagum pr. Alcochete”], becoming rounded and irregular in old trees. Foli-
A. X. Pereira Coutinho s.n. (holotype LISU). age branchlets numerous, patent or spreading with
a narrower angle, slender, quadrangular, ultimate
Juniperus navicularis Gand., Bull. Soc. Bot. Prance branchlets 1–1.5 mm diam., covered with gibbous
57: 55. 1910. scale leaves, persistent. Leaves decussate, sometimes
in alternate whorls of 3, closely appressed, scale-like,
0.5–1(–2) × 0.5–0.7 mm on (pen)ultimate branch-
Description
lets, broadly rhombic, gibbous; margins entire or
Shrubs to 2 m with fastigiate branching; foliage minutely denticulate; apex obtuse, sometimes acut-
branchlets spreading. Leaves spreading at 70–90 de- ish; stomata in two small groups abaxially near base,
grees from shoot, (4–)6–12 × 1.1.5 mm, ‘boat shaped’ adaxially in two narrow bands; glands conspicuous,
(but on whip shoots longer and linear-acicular), sometimes inconspicuous, central, oval to linear,
sub­obtuse to acute (pungent on whip shoots). Pollen usually active; leaf colour lustrous light green or
cones 2–3 mm long, ovoid. Seed cones subglobose to greyish green. Pollen cones numerous, terminal on
ovoid, 7–10 mm long, yellowish or reddish but not short ultimate branchlets, ovoid or short cylindrical,
glaucous or pruinose when growing, red when ripe. 2–4 mm long; microsporophylls 6–16, decussate or
sometimes ternate, peltate, with minutely denticu-
late margins, bearing 3–5 abaxial pollen sacs. Seed
Distribution
cones terminal on short branchlets, growing in two
SW Portugal, adjacent part of Spain. seasons to ochraceous, red-brown or purplish black,
TDWG codes: 12 POR SPA-SP sometimes pruinose, (sub)globose to ovoid, (5–)7–
12(–14) mm diam., soft pulpy or fibrous, maturing
soft succulent. Bract-scale complexes 4–6 (–9), dec-
Conservation
ussate or ternate, entirely fused, smooth, lustrous
IUCN: LC or slightly rugose when dry, with triangular bract
apices protruding from transverse ridges ­(sutures).
 Seeds 3–8 per cone, relatively small, unequal to Juniperus phoenicea L. subsp. phoenicea. Juniperus
4 mm long, triangular or flattened. phoenicea L. subsp. eu-mediterranea P. Lebreton
& S. Thivend, Naturalia Monspel., Bot. 47: 8. 1981
(nom. illeg., Art. 26). Type: [locality unknown
Distribution
(“Habitat in Europa australi, Monspeli”)], Herb.
Macaronesia: Canary Is., Madeira Is.; Mediterranean Burser XXV: 61 (lectotype UPS). Fig. 142, 143
Region: Albania, Algeria, Andorra, Bosnia-Herze-
govina, Bulgaria, Croatia, Cyprus, Egypt (Sinai), Juniperus canariensis Guyot & Mathou, Trav. Lab.
France, Greece, Italy, Jordan, Lebanon, Libya, Mo- Forest. Toulouse T. 1 (3, 20): 7. 1942; Juniperus tur-
rocco, Portugal, Romania, Spain, Tunisia, Turkey; binata Guss. subsp. canariensis (Guyot & Mathou)
434 Saudi Arabia (along Red Sea). Rivas-Martínez et al., Itinera Geobot. 7: 511. 1993.
TDWG codes: 12 BAL COR FRA-FR FRA-MO POR
SAR SPA-GI SPA-SP 13 ALB GRC ITA-IT KRI SIC-SI
Description
TUE YUG-BH YUG-CR 20 ALG LBY MOR-MO MOR-
SP TUN 21 CNY MDR 34 CYP EAI LBS-LB LBS-SY SIN Leaves small, gibbous, obtuse. Seed cones (sub)glo-
35 SAU bose, (5–)7–13 mm diam., smooth, lustrous, green-
ish or purplish green maturing to red-brown or
purplish black.
Ecology
In garrigue, maquis, or evergreen microphyllous
Distribution
woodland on dry, stony ground, limestone outcrops,
or sand dunes at altitudes between 1 m and 2400 m as for the species.
a.s.l. This species is commonly associated with Pinus
Conservation
halepensis, P. brutia, Quercus ilex, Pistacea lentiscus,
Cistus spp., Olea europaea, Lavandula spp., Artemi- IUCN: LC
sia herba-alba, and numerous other genera. There
is a predominance of limestone, but granitic rock, Juniperus phoenicea L. subsp. turbinata (Guss.)
sandstone, serpentine, volcanic rock, as well as sand Nyman, Consp. Fl. Europ. 3: 676. 1881. Juniperus
dunes are also mentioned as substrates. Soils are turbinata Guss., Fl. Sicula Syn. 2: 634. 1844.
usually rocky or skeletal and it can grow well from Type: [“In arenosis, vel rupestribus maritimis:
crevices in bedrock. The climate is Mediterranean, Montallegro; Secciara (Gasparr.)”], G. Gasparrini
with dry and hot summers. s.n. (FI?, PAV).

Uses Description
This species is rarely taken into cultivation in Medi- Leaves more elongate than in subsp. phoenicea, acut-
terranean countries and only a few cultivars have ish. Seed cones subglobose to ovoid, 12–14 × 9–11
been named of it. The wood is used in Algeria and mm, rugose, ochraceous or yellowish brown when
Tunisia for construction, fence posts and firewood; full grown.
in most other areas where it occurs it is of little eco-
nomic value. The reddish and more or less succulent
Distribution
cones (‘berries’) can be used in cooking and alco-
holic beverages. The subspecies turbinata forms a Mediterranean: France (Corse and mainland coast),
dense, prostrate shrub suitable in rock gardens and Greece, Italy (Sardinia, Sicily), Morocco, Portugal
as a ground cover in regions where it is not likely to (Algarve), Spain (Baleares and mainland coast),
be in danger of damage by frost. ­Tunisia.
TDWG codes: 12 BAL COR FRA-FR POR SAR SPA-
2 subspecies are recognized: SP 13 GRC SIC-SI 20 MOR-MO TUN
 decurrent, rhombic to broadly rhombic; margins
Ecology
minutely denticulate; apex appressed or sometimes
This subspecies is restricted to littoral maritime hab- free, acute/acuminate; stomata on abaxial side lim-
itats, on rocks or sand dunes. ited to decurrent leaf base, on adaxial surface in two
tapering bands; glands more or less conspicuous,
large, circular to elliptical, exudate often present;
Conservation
leaf colour yellowish green or light green. Pollen
IUCN: LC cones terminal, solitary, ellipsoid or ovoid-oblong,
3–4 × 2–2.5 mm; microsporophylls 8–10, decussate,
peltate, acute, upper margins minutely denticulate,
Juniperus pinchotii Sudw., Forestry & Irrig. 11: with 3–4 abaxial pollen sacs. Seed cones terminal on 435
204. 1905. Juniperus monosperma (Engelm.) Sarg. straight, very short branchlets, maturing to reddish
var. pinchotii (Sudw.) Melle, Phytologia 4: 29. brown or copper-brown with glaucous white bloom
1952. Type: USA: Texas, Brewster Co., Big Bend in 1–2 seasons, subglobose to globose, (5–)6–8(–10)
N. P., Palo Duro Canyon [“Palodura Canyon”], mm diam., internally soft pulpy, succulent. Bract-
G. L. Clothier s.n. (holotype US). scale complexes 4–6, entirely fused, decussate, with
upper pairs indiscernible in mature cones; bract api-
Juniperus erythrocarpa Cory, Rhodora 38: 186. 1936; ces (prominently) exserted; surface smooth, rugose
Juniperus pinchotii Sudw. var. erythrocarpa (Cory) when dry, glaucous. Seeds 1–2 per cone, broadly
Silba, Phytologia Mem. 7: 35. 1984. ovoid to pyriform, 4–6 × 3–4.5 mm; base truncate;
Juniperus texensis Melle, Phytologia 4: 26. 1952. apex acutish, smooth or shallowly grooved and
ridged on sides or with two opposite ridges, with
resin in the grooves or pits, light yellowish brown to
Etymology
chestnut brown, with a large, lighter bilobed hilum.
This species was named after Gifford Pinchot, Chief
Forester under US President Theodore Rooseveldt
Distribution
and an early conservationist.
USA: Arizona, New Mexico, Oklahoma, Texas;
Mexico: Chihuahua, Coahuila, Durango, Nuevo
Vernacular names
León, Sonora, Tamaulipas, Zacatecas.
Pinchot juniper, Redberry juniper TDWG codes: 74 OKL 76 ARI 77 NWM TEX 79
MXE-CO MXE-CU MXE-DU MXE-NL MXE-TA MXE-
ZA MXN-SO
Description
Large shrubs to 6 m, dioecious, rarely monoe-
Ecology
cious; multistemmed or rarely with a short trunk
and a maximal diam. of 20 cm, readily regenerating Juniperus pinchotii occurs in (semidesert) grassland
from burnt or cut stumps (coppicing). Bark of thick (“Bouteloua grasslands”) and in Pinyon-Juniper
branches and stems thin, slowly exfoliating in scales woodland with Pinus cembroides, locally P. pinceana
and strips; inner bark cinnamon; outer bark weath- or P. nelsonii, Juniperus spp., Quercus spp., Opuntia
ering ash-grey. Branches numerous, ascending from spp., and Yucca spp.; on escarpments or plateaus of
base or nearly erect, often curved, forming a broad, limestone or volcanic rock, or along sandy washes
rounded or more irregular and open crown. Foliage and on river terraces in sandy or gravely soil. The
branches numerous, irregularly disposed but not altitudinal range is 600–2100 m a.s.l. The climate is
pendulous, ultimate branchlets slender and stiff, up warm to cool temperate, with long dry spells and
to 25 mm long, 1.1–1.8 mm thick, subterete or weakly mostly summer rain coming as scattered storms.
quadrangular in cross section, covered with scale
leaves, persistent. Leaves in alternate whorls of 3, or
Conservation
decussate on ultimate branchlets, imbricate, scale-
like, 1–2(–3) × 0.5–1.5 mm on lateral branchlets, IUCN: LC
 2–3 abaxial globose pollen sacs. Seed cones axillary
Uses
on very short dwarfed shoots, growing to globose
No uses have been recorded of this shrubby juniper. or ovoid cones (5–)6–9 × 5–6 mm in two seasons,
It is rare in cultivation and would require open spac- ripening to lustrous purplish black or bluish black.
es with much of sunshine, while it can withstand oc- Bract-scale complexes 3, rarely 4, entirely fused, not
casional frost well. discernible in soft succulent-pulpy mature cones ex-
cept for the presence of 3(–4) subapical minute bract
tips. Seeds ovoid-globose, 5–6 × 4 mm, with several
Juniperus pingii W. C. Cheng, Bull. Soc. Hist. Nat. shallow, resinous grooves or pits towards base.
Toulouse 79: 76. 1944.
436
Distribution
Etymology
China: Gansu (Min Shan), Hubei (Daba Shan),
This species was named after Prof. C. Ping, founder S Qinghai, Shaanxi (Qin Ling Mts.), W Sichuan,
of the Biological Laboratory of the Science Society NW Yunnan, Xizang [Tibet].
of China. TDWG codes: 36 CHC-HU CHC-SC CHC-YN CHN-
GS CHN-SA CHQ CHT
Vernacular names
Ecology
Ping’s juniper; chui zhi xiang bai, xiang bai (var.
wilsonii) (Chinese) This species occurs from subalpine coniferous for-
ests with Abies, Larix, Picea and Pinus to beyond the
tree line in alpine meadows and high steppes to near
Description
the line of perennial snow or glaciers, at altitudes be-
Procumbent or erect shrubs to (small) trees, mon- tween 2650 m and 4850 m a.s.l.
oecious, to 10 m but reputedly attaining 30 m. Bark
on large stems fissured, exfoliating in long strips,
Uses
weathering grey-brown. Branches spreading or as-
cending, very dense in procumbent shrubs, form- The subalpine to alpine decumbent shrub forms are
ing a spreading or rounded crown. Foliage branches excellent plants for cultivation in cool temperate to
short, patent, drooping or downcurved, or pendu- cold climate regions. A few clones are in cultiva-
lous, ultimate branchlets in shrubs 10–30 mm long, tion in the West but are rare; these are commonly
in trees with pendulous branchlets much longer, J. pingii var. wilsonii. The tree form (var. pingii) is
stout or slender, subterete to more or less 6-angled, occasionally seen in cultivation in China; in Europe
sometimes quadrangular, covered with imbricate some introductions are likely to masquerade under
scale leaves. Leaves in alternate whorls of 3, rarely the names J. recurva and/or J. squamata which are
decussate, imbricate, decurrent at base, appressed actually this species. Several selected cultivars are in
with incurved apices or sometimes curved but free, the horticultural trade.
lanceolate (naviculate) to broadly subulate, free por-
tion broadest below middle section, incurved, (2–)3– 4 varieties are recognized:
5(–7) × 1–1.5 mm, concavo-convex; abaxial surface
distinctly keeled or ridged, especially towards the Juniperus pingii W. C. Cheng var. pingii. Type:
apex, smooth, glaucous green; adaxial surface with Illustration in Y. de Ferré, Bull. Soc. Hist. Nat.
two whitish bands proximally separated by a flat or Toulouse 79: 75. 1944 (lectotype).
slightly raised midrib; margins entire; apex acumi-
nate to pungent; epistomatic, stomata in two con-
Description
verging bands. Pollen cones axillary on very short
dwarfed shoots, solitary, globose, 3–4 mm diam.; Arborescent shrubs to trees possibly to 30 m tall.
microsporophylls 6–9, ternate, rarely decussate, Foliage branches drooping or pendulous; ultimate
more or less cordate, obtuse or cuspidate, bearing
branchlets usually 6-angled, or subterete, slender; Juniperus pingii W. C. Cheng var. miehei Farjon,
leaves nearly straight on young trees, later curved. Monogr. Cupressaceae & Sciadopitys: 346. 2005.
Type: China: Xizang (Tibet), Zangbo River, Upper
Zangbo basin, S of Saga, along road to Gyirong,
Distribution
G. Miehe & S. Miehe 9532-01 (holotype K).
China: W Sichuan, NW Yunnan, SE Xizang [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHT
Description
Procumbent shrubs with very dense, short foliage.
Conservation
Ultimate branchlets predominantly quadrangular,
The tree form is apparently quite rare, but its remote- with decussate, short leaves. Seed cones globose, 5–7 437
ness has until the present been a safeguard against mm diam.
threats from human interference.
IUCN: NT
Distribution
Juniperus pingii W. C. Cheng var. chengii (L. K. Fu China: Xizang [Tibet], known only from a locality
& Y. F. Yu) Farjon, Monogr. Cupressaceae & Scia- in the upper Zangbo River basin.
dopitys: 344. 2005. Juniperus chengii L. K. Fu & TDWG codes: 36 CHT
Y. F. Yu, Novon 7 (4): 443. 1998. Type: China: Yun-
nan, NW Yunnan, Zhongdian, (mountains near
Ecology
Zhongdian), T. T. Yü 10960 (holotype PE).
Found in juniper dwarf scrub patches in very open
alpine steppe on the Tibetan Plateau at 4850 m a.s.l.
Description
in the rain/snow shadow of the main crest of the
Small trees 3–9 m tall. Foliage branchlets slender, Himalayas.
2–3 mm thick, quadrangular to 6-angled, mostly
curved. Leaves decussate or in whorls of three, im-
Conservation
bricate, scale-like or nearly so, 1.5–2.5 mm long and
ca. 1 mm wide on ultimate branchlets (up to 5 mm IUCN: DD
long on older leafy branchlets), thick, more or less
keeled, incurved, with acuminate apex or obtuse. Juniperus pingii W. C. Cheng var. wilsonii (Rehd.)
Silba, Phytologia Mem. 7: 36. 1984. Juniperus
squamata Buch.-Ham. ex D. Don f. wilsonii Rehd.,
Distribution
J. Arnold Arbor. 1: 191. 1920. Type: China: Sichuan,
China: NW Yunnan (imperfectly known from the W Sichuan, E. H. Wilson 985 (lectotype E).
type collection near Zhongdian only).
TDWG codes: 36 CHC-YN Juniperus pingii W. C. Cheng var. carinata Y. F. Yu
& L. K. Fu, Novon 7: 443. 1998; Juniperus carinata
(Y. F. Yu & L. K. Fu) R. P. Adams, Biochem. Syst.
Conservation
Ecol. 28: 541. 2000.
The distribution of this form seems to be much more
limited than that of the species as a whole, but its ex-
Description
tent remains insufficiently known. It was described
as a small tree; the most common and widespread Procumbent to erect shrubs or arborescent shrubs
variety, J. pingii var. wilsonii (Rehd.) Silba is a low to 6 m tall. Foliage branches not pendulous, usually
shrub. curved, stout, subterete or prominently 6-angled.
IUCN: DD Leaves appressed or sometimes curved but free,
slightly to strongly incurved.
 Juniperus procera Hochst. ex Endl., Syn. Conif.: 26.
Taxonomic notes
1847. Type: Ethiopia: Gonder Prov., Amhara, Adda
The distinction between Juniperus pingii var. wilso- Mariam, near Enschedcap [“ad ecclesiam Adda
nii and Juniperus squamata has been a contentious Mariam prope Enschedcap”], G. H. W. Schimper 537
subject for some time. Rehder (op. cit.) originally (lectotype L).
described this taxon as only a form of J. squamata,
but it was treated as a variety of J. squamata in sub-
Etymology
sequent arboricultural and floristic literature, and
as a cultivar name under J. squamata in the fourth The species epithet means slender or tall.
edition of Dallimore’ & Jackson’s Handbook of Con-
438 iferae and Ginkgo (4th ed., 1966). It was transferred
Vernacular names
to Sabina pingii var. wilsonii in FRPS 7 (Cheng &
Fu, 1978); this became Juniperus pingii var. wilsonii African pencil cedar, African juniper, East African
in Flora of China 4 (1999) and again Sabina pingii cedar, Cedar; Arar (Arabian); Birbirssa, Tedh (Ethi-
var. wilsonii in Higher Plants of China 3 (Fu et al., opia); Deyib (Somalia); there are many local names
2000). Its morphological differences with J. squama- used in Kenya and Tanzania.
ta are mainly in the leaves, which are more similar to
those of J. recurva. In high altitude subalpine mead-
Description
ows (4000–4300 m) of Baima Shan, NW Yunnan
J. pingii var. wilsonii and J. squamata grow together, Trees to 30–40 m tall, monoecious or dioecious;
sometimes accompanied by a form that could be a trunk to 1.5–2 m d.b.h. Bark on larger stems fis-
hybrid of the two (pers. obs., September 2000). Oth- sured, exfoliating in long, narrow, fibrous strips,
erwise the two taxa are distinct in this area. (pale) brown weathering grey-brown. Branches
long, spreading or ascending, foliage branches
drooping or pendulous, lax, forming a pyramidal
Distribution
crown in young trees, but later broader, rounded
China: S Gansu, NW Hubei, S Qinghai, S Shaanxi, and finally flat-topped or open and irregular in old
Sichuan, NW Yunnan, Xizang [Tibet]. trees. Foliage branchlets ultimately slender, (weakly)
TDWG codes: 36 CHC-HU CHC-SC CHC-YN CHN- quadrangular, 0.6–1 mm wide, covered with close-
GS CHN-SA CHQ CHT ly appressed leaves, persistent. Leaves on mature
plants scale-like, decussate, imbricate, decurrent, on
ultimate branchlets 0.5–1 × 0.4–0.6 mm, triangular-
Ecology
rhombic, acute; margins entire; stomata in 2 incon-
A high altitude variety commonly replacing sub- spicuous lines on each side mostly near base; leaves
alpine coniferous forest or woodland above the tree abaxially glandular; gland central, conspicuous, el-
line, forming thickets or patchy mats of decumbent liptic or nearly linear; leaf colour yellowish green or
juniper among scree, rocks, or in alpine grassland or light green. Pollen cones numerous, solitary, termi-
steppe. The altitudinal range is 2650–4800 m a.s.l. nal on short branchlets, subglobose to ovoid, 3–5 ×
2–3 mm; microsporophylls 10–12, decussate, peltate,
with minutely denticulate margins, bearing 2–3
Conservation
abaxial pollen sacs near lower margin. Seed cones
IUCN: LC terminal on short, erect branchlets, maturing in
the second season to (sub)globose, 3–7 mm diam.,
brown, blue or purplish black, usually glaucous or
pruinose. Bract-scale complexes 4(–6), decussate,
entirely fused, bract tip small, ca. 0.3 mm, scale tis-
sue dry pulpy. Seeds (1–)2–3(–4) per cone, angular-
ovoid, more or less flattened on one side, 4–5 × 3–3.5
mm, yellowish brown, with lighter hilum at base.
 in Africa and India, but only on a limited scale; in
Distribution
horticulture it is mostly confined to public parks in
NE, E & S Tropical Africa: Congo Republic, Djibou- Ethiopia and Kenya, including cemeteries. Outside
ti, Eritrea, Ethiopia, Kenya, Malawi, Somalia, Sudan Africa it is only planted in a few botanic gardens;
(near Red Sea), Tanzania, Uganda, NE Zimbabwe; under glass in temperate climate regions or outside
Arabian Peninsula: Saudi Arabia (Asir Range), in warmer countries.
­Yemen.
TDWG codes: 23 ZAI 24 DJI ERI ETH SOM SUD 25
KEN TAN UGA 26 MLW ZIM 35 SAU YEM-NY Juniperus procumbens (Siebold ex Endl.) Miq.,
in Siebold & Zuccarini, Fl. Japon. 2: 59, t. 127, f.
3. 1870. Juniperus chinensis L. var. procumbens 439
Ecology
Siebold ex Endl., Syn. Conif.: 21. 1847. Type: Japan:
This species is forming evergreen Afromontane for- locality unknown [“Japonia”], P. F. von Siebold s.n.
est (also locally invading onto savanna where fires (lectotype K).
permit), either with pure stands of Juniperus procera,
or mixed coniferous, with Afrocarpus gracilior, Podo­
Etymology
carpus milanjianus, or conifer-mixed angiosperm,
with Olea chrysophylla, O. hochstetterii, Faurea The species epithet means ‘laying down’ and refers
saligna, Dombeya mastersii, Olinia rochetiana, Ilex to the growth habit.
mitis, Vepris nobilis and numerous smaller trees and
shrubs, e.g. Agarista salicifolia, Catha edulis, ­Buddleja
Vernacular names
spp., Cadia purpurea, Cussonia spicata, Dodonaea sp.,
Erica arborea, Euclea schimperi, Faurea sp., Maytenus Creeping juniper; Hai-byakushin, Sonare (Japanese)
spp., Nuxia congesta, and Olea spp. Pure stands are
usually evidence of establishment after forest distur-
Description
bance in the past. Juniperus procera occurs on moun-
tain slopes, summits, on escarpments and outcrops Decumbent shrub to 50 cm tall, creeping with long,
and in forested ravines in sand, loam or clay over prostrate stems to 5 cm diam., monoecious, some-
various rock types, e.g. basalt, volcanic ash and cin- times dioecious. Bark thin, scaly, exfoliating with
ders, granite, limestone, or metamorphic rock. The thin, elongated flakes or strips, brown. Branches
altitudinal range is 1370–3000 m a.s.l. The climate is spreading, whip shoots ascending, covering rocks
tropical montane, with a prolonged dry season. or soil. Foliage branches short, stiff, spreading in a
forward direction from main stems, young shoots
(leaves) glaucous, smaller branchlets spreading at
Conservation
30–50 degrees from axis, covered with forward-­
This species has been logged in many areas, but it is oriented leaves, persistent. Leaves in alternate
still too common to be threatened with extinction. whorls of 3, decurrent, with free part spreading at
Depletion of old growth forest groves of this spe- a narrow angle, on (pen)ultimate branchlets acicu-
cies threatens to occur in Kenya and Ethiopia and lar, rarely partly scale-like, convex with a median
from this ecological point of view there is certainly groove extending two third of leaf length abaxially,
a conservation issue regarding the only juniper in 4–10 mm long, ca. 1 mm wide near base, concave
subsaharan Africa. with a raised midrib adaxially, gradually tapering
IUCN: NT to an acuminate-pungent apex; stomata adaxially in
two merging bands bordered by green leaf margins,
abaxially on short decurrent leaves in two lateral
Uses
spots near the base, in long decurrent leaves in two
The larger trees of this species are prized for timber, lateral grooves; leaf colour glaucous green or bluish
having good, workable and decay-resistant wood. green with white stomatal areas. Pollen cones termi-
It is used for fence posts and shingles on roofs, for nal, solitary on dwarfed axillary shoots, subglobose,
construction, furniture, cabinet making, and the 3–4 mm diam.; microsporophylls 9–12, ternate, with
manufacture of pencils. It is grown in plantations entire, hyaline margins and obtuse apex, bearing
 3 abaxial pollen sacs. Seed cones terminal, solitary on (1839–1888), who explored Central Asia and (great-
dwarfed axillary shoots, growing irregularly ovoid er) Mongolia.
but usually subglobose when fully mature, 7–9 mm
diam., purplish blue or blackish blue. Bract-scale
Vernacular names
complexes 3, whorled, of equal size; bract tips barely
excerted below the apex of the cone; outer surface Przewalski juniper; qi lian yuan bai (Chinese)
smooth or rugose, inner tissue soft pulpy. Seeds (1–)
2–3 per cone, ovoid-globose, 3–5 mm long, more or
Description
less flattened except when single, light brown.
Trees, rarely shrubs, to 15(–20) m, monoecious; trunk
440 up to 1 m d.b.h. Bark on larger stems exfoliating in
Distribution
longitudinal strips, weathering whitish grey or grey-
Japan: Kyushu, Nansei-Shoto (Ryukyu Islands) brown. Branches spreading, smaller branches droop-
(Okinawa Group, Sakishima Group). ing to subpendulous, forming a broadly rounded or
TDWG codes: 38 JAP-KY NNS OGA irregular crown. Foliage branchlets slightly curved
or drooping to pendulous, quadrangular or sub-
terete, 1.2–1.5 mm wide if covered with appressed
Ecology
scale leaves, often with transitional leaves or juvenile
The ecology of this species is poorly known. It oc- leaves, appearing ‘ragged’, persistent. Leaves on ma-
curs at high altitudes near or on island mountain ture plants scale-like as well as acicular (scale leaves
summits [Ohwi, Fl. Japan: 118 (1965) has it on “sea- predominant on old trees), decussate, or 3-whorled,
shores [of] Kyushu” but this is J. rigida subsp. con- imbricate, decurrent, scale leaves on ultimate branch-
ferta], where it forms decumbent mats over rocks. lets 1.5–2.5(–3) × 1–1.5 mm, triangular-­rhombic, with
free apex, acute, transitional and juvenile leaves with
Conservation spreading parts 4–8(–10) × 1.5–1.7 mm; margins en-
tire, pungent; scale leaves weakly amphistomatic,
IUCN: LC
juvenile and transitional leaves epistomatic; stomata
in two bands separated by an inconspicuous midrib;
Uses
scale leaves abaxially glandular; gland near base, el-
This species is popular in gardens as a ground cover- liptic to orbicular, convex; leaf colour green or glau-
ing evergreen conifer shrub and a limited number cous green. Pollen cones solitary, terminal on short
of cultivars are in the trade. It is used more often branchlets, subglobose to ovoid-globose, 2–3 mm
in Japan than in Europe, although it is not uncom- long; microsporo­phylls 6–8(–10), decussate, peltate-
mon there. Unlike some other species with a similar cordate, with entire hyaline margins, bearing 2–3
growth habit, Creeping juniper ‘faithfully’ follows abaxial pollen sacs near lower margin. Seed cones
the contours of rock gardens without erecting itself terminal on short erect branchlets, maturing in the
or massing in one spot. It is therefore very effective second season to become ovoid, (8–)10–13 × 9–10
in such gardens, especially in traditional oriental mm, lustrous blue-black or purplish black, slightly
gardens. pruinose and soft. Bract-scale complexes 6, decus-
sate or (2×) 3-whorled, entirely fused; bract tips usu-
ally hidden or minute; surface smooth; scale tissue
Juniperus przewalskii Kom., Bot. Mater. Gerb. succulent, resinous. Seeds 1(–2) per cone, ovoid-glo-
Glavn. Bot. Sada RSFSR 5: 28. 1924. Type: China: bose, laterally compressed or slightly flattened, (7–)
Gansu, Qilian Shan, Daban Shan, [“jugum a fluv. 8–12 × (6–)7–10 mm, shallowly or deeply grooved,
Tetung versus in regione alpina sol. humido, 13.000 with resin pits, light brown.
ft. alt.”], N. M. Przewalski 318 (lectotype LE).
Distribution
Etymology
China: Gansu, E Qinghai, N Sichuan (upper tribu-
This species name commemorates the Polish sol- taries of the Huang He).
dier, traveller and naturalist Nikolai M. Przewalski TDWG codes: 36 CHC-SC CHN-GS CHQ
 Ecology Description
Juniperus przewalskii occurs in the ecotone between (Decumbent) shrubs or small trees to 8–10 m, dio-
subalpine forest to alpine meadows or steppe, as well ecious, sometimes monoecious; multistemmed or
as in Juniper-Picea crassifolia woodland, on steep, monopodial, with a short trunk up to 1 m d.b.h. Bark
usually S-facing, calcareous slopes, with ground on larger stems shaggy, exfoliating in short strips
cover of grasses (e.g. Stipa splendens), shrubs (e.g. remaining partly attached, weathering grey-brown.
Salix sp.) or perennial herbs (e.g. Polygonum vivi- Branches ascending, assurgent or spreading, smaller
parum), on dry or moist substrates. The altitude branches numerous, short, stiff and spreading, form-
ranges between 2250 m and 3900 m a.s.l. The climate ing a dense, spreading or irregular crown. Foliage
is cold temperate with substantial periods of snow branchlets thick, quadrangular, sometimes more or 441
cover in winter. less terete, 1.5–2 mm wide, covered with appressed
or sometimes apically free leaves, persistent. Leaves
on mature plants normally scale-like, decussate,
Conservation
imbricate, decurrent, 1.3–2 × 1–1.2 mm, triangular-
The relatively restricted distribution and its occur- rhombic, gibbous, obtuse, on older branchlets to 4 ×
rence in a region where forest cover is scattered and 1.5 mm, mostly appressed but several with free apex;
limited to suitable slopes make it potentially vulner- margins entire or sometimes minutely and intermit-
able to logging and/or deforestation resulting from tently denticulate; stomata in 2 conspicuous short
increased human pressure on these resources. As bands on each side mostly near base; scale leaves
yet, however, there is no immediate concern regard- abaxially glandular; gland central in a groove, ellip-
ing this species. tic or oblong, sometimes absent; leaf colour green or
IUCN: NT yellowish green, sometimes slightly glaucous green.
Pollen cones numerous, solitary, terminal on short
branchlets, subglobose to globose, 2–3 mm long;
Uses
microsporophylls 6–8, decussate, peltate-cordate,
No uses have been recorded, but this species is likely with rounded entire hyaline margins, bearing 2–3
to be used for firewood as it is one of the few trees abaxial pollen sacs. Seed cones terminal on short
that exists in some parts of its range. erect branchlets, maturing in the second season to
become ovoid, (7–)8–14 × (6–)7–10 mm, lustrous
blue-black or purplish black and soft. Bract-scale
Juniperus pseudosabina Fisch. & C. A. Mey., Index complexes 4–6, decussate, entirely fused; bract tip
Sem. Hort. Petrop. 8 (1841): 24, 65. 1842. Type: ca. 0.5 mm; surface smooth; scale tissue succulent,
Kazakhstan: Dzhungarskiy Alatau, Tarbagatay Mts., resinous. Seeds 1 per cone, ovoid-ellipsoid, laterally
A. G. von Schrenk s.n. (lectotype K). Fig. 144, 145 compressed, 6–8 × 4.5–6.5 mm, shallowly grooved,
light brown.
Juniperus turkestanica Kom., Bot. Mater. Gerb.
Glavn. Bot. Sada RSFSR 5: 26. 1924; Juniperus
Taxonomic notes
pseudo­sabina Fisch. & C. A. Mey. var. turkestanica
(Kom.) Silba, Phytologia Mem. 7: 36. 1984. Russian botanists and ecologists treat the taller
shrub in the west as a distinct species J. turkestanica
Kom. No botanical characters appear to separate the
Etymology
two and J. turkestanica is not a species, but only an
The species epithet means ‘similar but not equal to ecotype that predominates in the juniper forest and
(J.) sabina’, a species that occurs in the same region. woodland, an ecosystem not present in the eastern
parts of the Central Asian mountains.
Vernacular names
Distribution
xin jiang fang zhi bai, ka shi fang zhi bai, kun lun
fang zhi bai (Chinese); no local (Kyrgiz) names have Central Asia: Afghanistan (Takhar), China
been recorded. (Xinjiang), Kazakhstan (southern mountains),
 ­ irgyzstan, Mongolia, Pakistan (Baltistan, Hindu
K Juniperus recurva Buch.-Ham. ex D. Don, Prodr.
Kush, Karakoram Range), Tadjikistan, Uzbekistan Fl. Nepal. (2): 55. 1825.
(Turkestan Range).
TDWG codes: 32 KAZ KGZ TZK UZB 34 AFG 36
Etymology
CHX 37 MON 40 PAK
The species epithet refers to the (re)curved adult
type leaves.
Ecology
Occurring in subalpine conifer forest with Picea
Vernacular names
schrenkiana, Pinus sibirica, or P. wallichiana, in ju-
442 niper woodland (J. semiglobosa) and in montane Drooping juniper, Sacred juniper, Coffin juniper
to subalpine scrubland and steppe (predominant- (var. coxii); Pama (Hindi); chui zhi bai, xiao guo
ly Seriphidium maritimum [Artemisia maritima] chui zhi bai (var. coxii) (Chinese)
steppe rich in grasses and geophytes), with e.g.
J. sabina, Cotoneaster, Kobresia capillifolia, Rhodo-
Description
dendron anthopogon, Rosa, and Salix. The altitudi-
nal range is from 1950 m to 4100 m a.s.l. It is often Shrubs or trees to 15(–40) m tall, monoecious, oc-
restricted to rocky outcrops and S-facing slopes, es- casionally dioecious; multistemmed or monopodial;
pecially in forests, and occurs on various rock types trunk up to 2 m d.b.h. Bark on larger stems exfo-
and soil types from coarse gravel terraces to dry S- liating in longitudinal, curling strips, red-brown
exposed loess slopes. The climate is extreme conti- weathering grey-brown. Branches ascending or
nental with short, hot, dry summers and long, cold, spreading, higher order branches assurgent and
snowy winters. Both climatic factors and (possibly) drooping or very long and pendulous, forming a
grazing pressures are responsible for a shift from broad pyramidal to eventually rounded or irregular
erect to decumbent shrubs from west to east in its crown. Foliage branchlets numerous, crowded, lax,
range. more or less terete, covered with incurved or slightly
spreading leaves, persistent. Leaves in alternating
whorls of 3 (on whip shoots occasionally decussate),
Conservation
imbricate, decurrent, mature type leaves 2–10 × 0.8–
IUCN: LC 1(–1.2) mm, lanceolate-acicular, weakly keeled; mar-
gins entire, acute to pungent or acuminate, incurved
or straight with a free apex; stomata abaxially in
Uses
2 inconspicuous lines on each side mostly near base,
This species is more often shrubby than a tree and in two adaxial white primary bands separated by an
consequently it is less often used for firewood or obscure midrib; leaf colour green or olive-green.
small timber. The large blue cones (‘berries’) are soft Intermediate leaf shapes, as well as juvenile-type
and probably edible, but no commercial use of them leaves, usually present in mature trees and retained
was spotted in local markets on a trip to Kirgyzstan in one variety. Pollen cones numerous, solitary, ax-
in August 2000. Especially the shrubby form of this illary on dwarfed shoots, ovoid-oblong, 5–6 × 2–3
species would make a very atractive juniper in cul- mm long; microsporophylls 10–16, decussate, more
tivation but it seems to be absent in gardens; this is or less remote, peltate-cordate, with entire-hyaline
an omission that ought to be rectified. The species to erose margins, bearing 3 abaxial pollen sacs. Seed
occurs in a continental climate at high elevations in cones axillary on dwarfed shoots, maturing in the
mountains similar to the Alps and should be hardy second season becoming ovoid or subglobose, 6–12
even in high latitudes. × 5–9 mm, lustrous purplish black and soft. Bract-
scale complexes 3, entirely fused; bract tip subapical,
ca. 0.5 mm; surface smooth; scale tissue succulent,
resinous. Seeds 1 per cone, ovoid-globose, broadest
at base, 6–8 × 5–6 mm, shallowly grooved with dark
brown resin pits near base, light brown.
 6

4
443

3
1

9 7
10 8

Pl ate 18 . Juniperus recurva var. recurva. 1. Habit of tree. 2, 3. Branches with leaves and seed cones. 4.
Branchlet with leaves. 5, 6. Leaves. 7. Branchlet with pollen cone. 8. Microsporophyll with open pollen sacs
and pollen. 9. Seed cone. 10. Seed.
 Distribution Distribution
Himalaya: Nepal, Sikkim, Bhutan, India (Arunachal As for the species.
Pradesh, Assam); Myanmar [Burma]; China: S Xi-
zang [Tibet], NW Yunnan, SW Sichuan.
Ecology
TDWG codes: 36 CHC-SC CHC-YN CHT 40 ASS-AS
EHM-AP EHM-BH EHM-DJ EHM-SI NEP WHM-UT In high montane to subalpine coniferous forest, with
41 MYA Abies spp. or Picea spp. and an understorey of Rhodo-
dendron, in Rhododendron thickets in the ecotone be-
tween forest and alpine meadows, and in the latter in
Ecology
444 mixed shrub communities with e.g. Rhodo­dendron,
In montane evergreen rainforest (var. coxii) ascend- Salix, Cotoneaster, Berberis, ­Lonicera, Spiraea, and
ing to and beyond the tree line in alpine scrub and Potentilla. The altitudinal range is 2500–4500 m a.s.l.
rocky meadows (var. recurva). The altitudinal range It is common in rocky areas (e.g. moraines) or in al-
is 2400–3800 m a.s.l. See for more detailed accounts pine meadows strewn with boulders, usually on sili-
under the varieties. ceous rock. In western Yunnan I observed that this
variety is a pioneer after destructive forest fires that
had killed most trees of Abies and Tsuga in the area.
Uses
It was not clear if, but it could be possible that the
In its native countries this species is used for timber extensive stands of J. recurva var. recurva observed
as well as ornamental trees in gardens of monaster- in this area (Nu Shan, NW of Caojian) are the result
ies and temples. The wood and foliage are burned of past forest destruction. The climate is more alpine
for incense in Buddhist temples. In Myanmar [Bur- with much longer periods of deep snow cover with
ma] the wood of large trees is used to make coffins. this variety than with var. coxii.
The drooping form J. recurva var. coxii is a highly
ornamental tree much valued and often planted in
Conservation
regions of Europe with a mild, moist climate. Sev-
eral cultivars have been selected, some with even IUCN: LC
more pendulous foliage than var. coxii. Decumbent
shrubs in cultivation under the name J. recurva (un- Juniperus recurva Buch.-Ham. ex D. Don var. coxii
less cultivars from proven J. recurva provenance) (A. B. Jacks.) Melville, Bull. Misc. Inf. R. B. G. Kew
are probably J. pingii var. wilsonii; J. recurva is not 1958: 533. 1959. Juniperus coxii A. B. Jacks., New Fl.
a decumbent shrub in the wild, but is often multi- & Silva 5: 33. 1932. Type: United Kingdom: England,
stemmed. It is one of the few junipers that require cultivated in Exbury Gardens, Hampshire; from
a good amount of rainfall and thrive best in a mari- R. Farrer & E. H. M. Cox 1407 collected in Myanmar
time, cool but mild climate. (Burma), Chimili Valley (26.3 N 98.6 E) in 1920,
R. Farrer 1407 (holotype BM).
2 varieties are recognized:
Description
Juniperus recurva Buch.-Ham. ex D. Don var. re-
curva. Type: Nepal: Narainghet (?) [“Narainhetty”], Trees up to 40 m tall, 2 m d.b.h., monopodial; foliage
F. Buchanan-Hamilton s.n. (holotype BM). Pl. 18 branches long, pendulous. Juvenile or transition type
leaves retained, 6–10 × 0.8–1 mm, straight or slightly
incurved, spreading at 30–45 degrees from the shoot.
Description
Seed cones 6–8 × 5–6 mm; seeds 5–6 × 3–4 mm.
Shrubs or small trees to 15 m tall, foliage branches
drooping, not pendulous. Leaves 2–3(–5) × 1–1.2
Distribution
mm, sometimes a few branches with intermediate
leaves slightly longer, incurved. Seed cones 7–12 × China: NW Yunnan, SE Xizang [Tibet]; Bhutan;
6–9 mm; seeds 6–8 × 5–6 mm. N Myanmar [Burma]; Sikkim; India, Arunachal
Pradesh?
TDWG codes: 36 CHC-YN CHT 40 EHM-BH EHM- when covered in leaves, with leaf whorls dense (8–
DJ EHM-SI 41 MYA 10 per cm length of shoot) or sparse (2–3 per cm),
turning light brown to reddish brown. Leaves ter-
nate, in alternating whorls, spreading 40–90° from
Ecology
shoot axis, not decurrent, (7–)10–20(–23) × 1–1.7
In temperate montane evergreen rainforest, either mm, ­bifacial-triquetrous, narrowly linear-acicular,
coniferous forest, with Abies forrestii, A. densa, straight or slightly curved, abaxially ridged or nearly
A. delavayi, Larix griffithii, L. potaninii, Picea spinu- flat; margins entire; apex pungent; epistomatic, sto-
losa, P. likiangensis, and Pinus wallichiana, or mixed mata in a single, narrow white band often nearly ob-
forest, with e.g. Abies-Tsuga-Lauraceae-Fagaceae, of- scured in a narrow groove; leaf colour light green.
ten with an understorey of Rhododendron spp. The Pollen cones on dwarf shoots in axils of leaves, sub- 445
altitudinal range is 2400–3800 m a.s.l. In these forest globose, 3–5 × 3–4 mm; microsporophylls 9–12, ter-
types the juniper usually occupies rocky slopes or nate, peltate-cordate, with raised midrib and often
gullies, or disturbed and degraded sites where the caudate apex, bearing abaxially 4–6 globose yellow
canopy has been opened substantially due to felling pollen sacs. Seed cones on axillary, very small (1–2
and grazing. mm) dwarf shoots covered with whorled 1 mm long
scale leaves, (sub)globose to broadly ovoid, 5–8(–11)
× 5–6(–8) mm, maturing within 18 months, becom-
Conservation
ing smooth, soft pulpy, resinous, brown to bluish
Due to deforestation and logging in many areas of black, often pruinose. Bract-scale complexes 3(–6),
SW China that infringes on the forests in which J. re- ternate, of which only 3 develop to form the cone,
curva var. coxii occurs, its area of occupancy (AOO) completely fused with suture lines only visible at
has been reduced substantially and this form is now apex, smooth, becoming rugose in sicco. Seeds 3,
difficult to find in many areas of NW Yunnan. 4–5(–6) × 3–3.5(–4.5) mm, angular, slightly flattened
IUCN: VU (A1c) on one or two sides, ridged; apex obtuse or rounded;
brown with darker resin pits near the base.

Juniperus rigida Siebold & Zucc., Abh. Math.-Phys.

Distribution
Cl. Königl. Bayer. Akad. Wiss. 4 (3): 233. 1846.
China: Gansu, N Hebei, Heilongjiang, Jilin, Liaon-
ing, Nei Monggol [Inner Mongolia], Ningxia, E
Etymology
Qinghai, Shaanxi, Shanxi; Japan: Hokkaido, Honshu,
The species epithet means ‘rigid’, or ‘stiff ’ and per- Kyushu, Nansei-Shoto (Ryukyu Islands), Shikoku;
haps refers to the needle leaves. North Korea; Russian Far East: Primorje, Sakhalin.
TDWG codes: 31 PRM SAK 36 CHI-NM CHI-NX
CHM-HJ CHM-JL CHM-LN CHN-GS CHN-HB CHN-
Vernacular names
SA CHN-SX CHQ 38 JAP-HK JAP-HN JAP-KY JAP-SH
Needle juniper, Temple juniper; Nezu, Muro, Nijiko, NNS KOR-NK
Nezumisashi, Miyamanezu (Japanese)
Ecology
Description
From sandy ocean shores (subsp. conferta) to up-
Decumbent or erect shrubs, or small trees to 10 m land moors, grassy or shrubby mountain slopes and
tall, dioecious, multi-stemmed or monopodial, open woodland, on a variety of rocks and soils. Al-
d.b.h. up to 40–50 cm. Bark on trunks of trees fis- titudinal range 1–2200 m a.s.l. See for more detailed
sured, fibrous, exfoliating in thin, narrow strips, accounts under the subspecies.
grey-brown. Branches decumbent, spreading or
ascending, smaller branches in trees drooping or
Uses
pendulous, forming a conical or pyramidal crown.
Foliage branches dense and short erect or more re- This species, in particular its decumbent growth
mote and long pendulous, triangular in cross ­section forms, has been in cultivation since many centuries
 in Japanese gardens and in temple grounds. The de- Juniperus rigida Siebold & Zucc. subsp. conferta
cumbent subsp. conferta is used for bonsai culture. (Parl.) Kitam., Acta Phytotax. Geobot. 26 (1–2): 9.
It is much less commonly used elsewhere, but is very 1974. Juniperus conferta Parl., Conif. Nov.: 1. 1863;
suitable for ground cover planting in large rock gar- Juniperus rigida Siebold & Zucc. var. conferta (Parl.)
dens, although its needle leaves are very sharp. The Patschke, Bot. Jahrb. Syst. 48: 678. 1913. Type: Japan:
tree form (subsp. rigida) is commonly planted in [locality unknown, (probably Ryukyu Islands)],
gardens and in temple grounds in Japan, sometimes C. Wright s.n. (holotype K).
in Korea and China. The wood is of local impor-
tance only and used for furniture, wood carving and Juniperus rigida Siebold & Zucc. subsp. litoralis
other domestic uses. Urussov, Bjull. Glavn. Bot. Sada 122: 56. 1981; Juni-
446 perus rigida Siebold & Zucc. var. litoralis (Urussov)
2 subspecies are recognized: Kozhevnikova, Fl. Ross. Dal’nego Vostoka: 42. 2006.

Juniperus rigida Siebold & Zucc. subsp. rigida. Vernacular names

Type: Japan: Honshu [“in Japoniae ad monte
Shore juniper; Hai-nezu, Hai-muro, Hai-matzu
Hakone” (Hakonegasaki?)], J. Bürger, [comm.
(Japanese)
P. F. von Siebold 1842] s.n. (lectotype M).
Description
Description
Decumbent shrubs with assurgent or erect branch-
Erect shrubs or small trees, branches spreading. Fo- es. Foliage branchlets short, erect, densely set with
liage branchlets drooping to long pendulous; leaf leaves in whorls up to 8–10 per cm length of shoot.
whorls more or less remote with 2–3 whorls per cm Leaves 1–1.7 mm wide with a correspondingly wider
length of shoot; leaves 1–1.3 mm wide, with very nar- stomatal band. Seed cones subglobose to broadly
row stomatal band in a deep groove. Seed cones 5–8 ovoid 8–11 mm long. Seeds 5–6 × 4–4.5 mm.
× 5–8 mm; seeds 4–5 × 3–3.5 mm.
Taxonomic notes
Distribution
This decumbent juniper is sometimes still treated as
As for the species, but not on Sakhalin Island. a distinct species. Apart from the different growth
form, its leaves are larger (wider) than those in the
erect growing form of the species. Ontogenetically,
Ecology
this leads to larger ovuliferous cones and larger seeds
Juniperus rigida subsp. rigida is common in sec- as well. As with the Mediterranean coastal subspe-
ondary scrubland or grassland, or in open for- cies J. oxycedrus subsp. macrocarpa, J. rigida subsp.
est or woodland, sometimes on sparsely vegetated conferta is apparently a littoral variant that is ecolog-
sandhills. The altitudinal range is 10–2200 m a.s.l. ically separated from the remainder of the species.
In pine forests it occurs with Pinus densiflora or The two subspecies have quite similar morphologi-
P. koraiensis, also with Betula, Quercus, and Tilia cal traits, which are perhaps of adaptive value to the
amurensis; in the shrub layer with e.g. Lespedeza bi- shifting dune sands on maritime coasts.
color, Cotoneaster and Rosa. It grows equally well on
moorland, sand dunes, sand flats, rocky mountain
Distribution
slopes and exposed limestone clifs. The climate is
mesic with abundant rainfall throughout the year; Japan: Hokkaido, Honshu, Kyushu; Russian Far
its distribution is largely determined by edaphic East: Sakhalin.
conditions. TDWG codes: 31 SAK 38 JAP-HK JAP-HN JAP-KY

Conservation Ecology
IUCN: LC This subspecies is strictly littoral and occurs on
sandy ocean shores, forming dense, spreading mats
of shrubby vegetation covering old beaches and two small bands. Pollen cones terminal on ultimate
dunes. Usually the vegetation is very open and has branchlets with scale leaves, 3–4 mm long, ellipsoid;
a pioneer character, but this juniper will sometimes microsporophylls 10–16, decussate, peltate-cordate,
occur (persist) in Pinus woodland near the shore. convex; upper margin erose, abaxially bearing 2–4
oblong pollen sacs. Seed cones terminal on recurved
short shoots 3–10 mm long with small scale leaves,
Conservation
maturing in 1–2 years becoming (irregularly) glo-
IUCN: LC bose, 4–8 mm, usually soft pulpy, resinous, purplish
brown, black-brown or blue. Bract-scale complexes
4, decussate, entirely fused, sutures partly visible as
Juniperus sabina L., Sp. Pl. 2: 1039. 1753. a curved ridge terminating in a minute bract tip. 447
Seeds 1–3(–5) usually 2 per cone, ovoid-flattened,
3–5 mm long, with proximal, shallow resin pits and
Etymology
longitudinal, shallow grooves, yellowish brown.
The origin of this epithet is probably the French
name of this species.
Distribution
SW, Central & SE Europe, N Africa (Atlas Mts.),
Vernacular names
Ukraine: Krym [Crimea], Caucasus, Russia (Altai
Savin juniper; Savin, Sabine (French); Sabina (Span- & Ural Mountains), Iran, Kazakhstan, Kirgyzstan,
ish); Sadebaum, Sebenbaum, Stinkwacholder (Ger- Turkey, Mongolia, N & NW China.
man); kazakij mozzevel’nik (Russian) TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SK
GER POL SWI 12 FRA-FR SPA-SP 13 ALB BUL GRC
ITA-IT KRI ROM SIC-SI TUE 14 KRY RUE RUS UKR-
Description
MO 20 ALG 30 ALT TVA WSB 32 KAZ KGZ 33 NCS
Usually decumbent, sometimes ascending or erect TCS 34 IRN TUR 36 CHI-NM CHI-NX CHM-HJ CHN-
shrubs, rarely a stunted tree to 10–12 m with a short, GS CHN-SA CHQ CHX 37 MON
leaning trunk to 2 m diam., dioecious or monoe-
cious. Bark on branches smooth, soon flaking, yel-
Ecology
lowish brown, becoming grey-brown. Branches
spreading horizontally or assurgent, of higher order In montane to alpine vegetation formations, ranging
mostly assurgent or erect, extending ‘broom-like’, from alpine scree and rocks or meadows to nearly
forming matting crowns in decumbent shrubs and closed coniferous forest; in the east of its Asian
dense, flat crowns in taller plants and small trees. range also in steppes and deserts. Altitudinal range
Foliage branchlets numerous, assurgent to erect, 400–3350 m a.s.l. More detailed accounts are given
slender or sometimes thicker 0.8–1(–1.4) mm diam., under the varieties.
subterete to nearly quadrangular in branchlets with
scale leaves, persistent. Leaves decussate, ocassion-
Uses
ally in alternating whorls of 3, short decurrent, of
two types: needle-like and, more commonly, scale- The decumbent shrub form has long been in culti-
like (acicular leaves in young plants and persisting vation in Europe and is relatively common; several
in one variety). Needle-like leaves linear-subulate, cultivars (some with fastigiate growth habit) have
(4–)6–10 × 0.5–1 mm, pungent, with a single or two been named. Its cultivation is often more or less
adaxial stomatal bands wider than green margins. restricted to countries where it is also native and
Scale leaves always decussate, imbricate, closely ap- where growers have experimented with this stock
pressed, on ultimate branchlets 1–2.5 × 0.6–1 mm, to produce cultivars; in other countries forms of the
ovate-rhombic to rhombic-lanceolate, abaxially similar species J. chinensis seem to prevail. Some of
with a conspicuous central, elliptic, yellowish gland, the cultivars of J. sabina retain juvenile type (nee-
dark lustrous green to yellowish green, strongly aro- dle) leaves, most have predominantly or exclusively
matic when bruised; margins entire; apex obtuse scale leaves in mature plants. Forms with needle
to acute; epistomatic or amphistomatic, stomata in leaves also occur in nature, so selection of this trait
 for horticulture is very easy where this variety (var. accounts for its wider distribution in Asia into the
davurica) is available. The wood is of little value, but Artemisia steppe and desert zones (var. arenaria). It
was traditionally used in the Alps to make walking is often found on limestone substrates but occurs on
sticks. Oil is distilled from branches and foliage and granitic rock as well, especially on drier slopes.
used for medicinal purposes; it has powerful diu-
retic properties.
Conservation
3 varieties are recognized: IUCN: LC

Juniperus sabina L. var. sabina. Type: [locality un- Juniperus sabina L. var. arenaria (E. H. Wilson)
448 known (“Habitat in Italia, Sibiria, Olympo, Ararat, Farjon, Monogr. Cupressaceae & Sciadopitys:
Lusitania”)], Herb. Burser XXV: 59 (lectotype UPS). 366. 2005. Juniperus chinensis L. var. arenaria
Fig. 146 E. H. Wilson, J. Arnold Arbor. 9: 20. 1928; Juniperus
arenaria (E. H. Wilson) Florin, Acta Horti Berg. 14
Sabina vulgaris Antoine var. yulinensis T. C. Chang (8): 353. 1948. Type: China: Qinghai, Qinghai Lake,
& C. G. Chen, Acta Phytotax. Sin. 19 (2): 263. 1981; J. F. Rock 13346 (holotype A).
Juniperus sabina L. var. yulinensis (T. C. Chang &
C. G. Chen) Y. F. Yu & L. K. Fu, Novon 7 (4): 444. Juniperus sabina L. var. mongolensis R. P. Adams,
1998. Phytologia 88 (2): 182. 2006.
Juniperus sabina L. var. monosperma C. Y. Yang, Fl.
Xinjiangensis 1: 305. 1993.
Description
Decumbent or spreading shrubs to 1.5 m. Foliage
Description
dense, assurgent or ascending, with short ultimate
Commonly decumbent shrubs, occasionally spread- branchlets, more or less glaucous. Leaves scale-like,
ing shrubs or a small, stunted tree. Branches as- 1.5–3 × 0.7–1 mm, appressed, obtuse, or longest (2–3
surgent, erect or spreading. Leaves of two types: mm) leaves with free apex, acute to pungent. Seed
acicular and scale-like; acicular leaves usually on cones more or less broad pyriform, 5–8 mm, with
seedlings and young plants to 10 years old, merging (1–)2–4(–5) seeds, yellowish glaucous.
to scale leaves on most mature plants. Seed cones
with (1–)2–3(–4) seeds, most frequently 2.
Distribution
China: N Gansu, Nei Mongol (I. Mongolia), Qing-
Distribution
hai, Shaanxi; Mongolia.
As for the species. TDWG codes: 36 CHI-NM CHN-GS CHN-SX CHQ 37
MON
Ecology
Ecology
In montane to subalpine coniferous forests of Larix,
Picea and Pinus, gradually replacing these where un- Artemisia steppes and grass steppes, sand dunes,
der human-imposed grazing regimes; also invading sparse vegetation on flood plains of scree and
into alpine meadows when old grazing patterns are gravel, S-facing mountain slopes, canyons and es-
changed, e.g. intensified. In Central Asian moun- carpments, forming patches of dwarfed scrub with
tains it occurs on S-facing slopes in mountain pas- herbaceous plants and grasses. The altitudinal range
tures in a characteristic pattern of rounded patches, is 2150–3350 m a.s.l.
often mixed with J. pseudosabina. Its altitudinal
range is 700–3000 m a.s.l. This subspecies is most
Conservation
abundant on sunny, dry slopes in mountains with
a mesic climate like the Alps; its drought tolerance IUCN: LC
figure 12 1. Falcati­
folium falciforme trees
in the Crocker Range,
Borneo

figure 123. Falcati­

folium falciforme seedling
at Fraser’s Hill, Malaysia

figure 12 6. Fitzroya cupressoides foliage

and seed cones

figure 122. Falcati­folium falciforme

flushing leaves

figure 12 4. Falcati-
folium taxoides on Mt.
Panié, New Caledonia

figure 12 5. Fitzroya
cupressoides in the N.P.
Alerce Andino (photo
P. Woltz)
 figure 12 8. Glyptostrobus pensilis pollen
cones and seed cones (photo D. White)

figure 127. Fokienia hodginsii

foliage and cones

figure 130. Halocarpus bidwillii foliage

f i gu re 1 29 . Halocarpus bidwillii in
North Island, New Zealand

fi g u re 13 1 . Halocarpus biformis foliage figure 132. Juniperus californica in Anza Borrego Desert State Park,
­California
figure 133. Juni-
perus californica seed
cones

figure 13 4. Juni­
perus chinensis var.
sargentii foliage and
seed cones
fig u re 135 . Juniperus communis var. fig ure 13 8. Juniperus deppeana var. figure 1 4 1. Juniperus oxycedrus subsp.
communis foliage and seed cones deppeana trunk with bark macrocarpa foliage and seed cones

f i gu re 136. Juniperus communis var. saxatilis

in Mt. Rainier N.P., Washington
f i gu re 139 . Juniperus flaccida var. flaccida
tree in Oaxaca, Mexico

figure 1 40. Juni-

perus occidentalis var.
australis tree in the
Sierra Nevada

figure 137. Juni-

perus deppeana var.
deppeana in Puebla,
Mexico
 f i g u re 1 4 2 . Juniperus phoenicea subsp. phoenicea figure 1 43. Juniperus phoenicea subsp.
at Cape St. Vincent, Portugal phoenicea foliage and cones

figure 1 45. Juniperus pseudosabina

seed cones in Kirgyzstan

fi g u re 1 4 4 . Juniperus pseudosabina in the Alaj Mountains, Kirgyzstan

figure 1 4 8. Juniperus semiglobosa

f i g u re 1 4 6 . Juniperus sabina var. sabina in Kirgyzstan foliage and seed cones

N.B. Figure 147 top left on facing page.

 figure 1 4 9. Juniperus semiglobosa pollen cones

figure 1 4 7. Juniperus
semiglobosa in Kirgyzstan

figure 15 1. Keteleeria
davidiana var. davidiana
foliage and seed cones

fig ure 150. Juniperus

virginiana var. virginiana
tree in North Carolina, USA

figure 152. Keteleeria

davidiana var. davidiana
seed cone

f i g u re 15 4 . Lagarostrobos franklinii
­foliage and pollen cones

f i g u re 15 3. Lagarostrobos franklinii tree

at Riveaux Creek, Tasmania
 figure 156. Larix decidua figure 15 7. Larix decidua var.
var. decidua bark decidua seed cones

f i gu re 15 5 . Larix decidua var. decidua

in the Alps, Switzerland

figure 159. Larix griffithii var. griffithii

seed cone

figure 161. Larix

lyallii in the Wenatchee
Mts., Washington, USA

f i g u re 15 8 . Larix gmelinii var.

principis-rupprechtii seed cones

f i gu re 16 0 . Larix kaempferi seed cones

 figure 164. Libocedrus bidwillii
foliage and seed cones

f i gu re 162. Lepidothamnus fonkii in Chile (photo M. Gardner)

figure 165. Libocedrus plumosa foliage

f i gu re 163 . Libocedrus bidwillii in North Island, New Zealand

fi g u re 166 . Manoao colensoi f i g ure 167. Metasequoia

seed cones (photo B. P. J. Molloy) ­glyptostroboides bark

figure 168. Metasequoia glyptostroboides

seed cones

figure 16 9. Microbiota decussata foliage

 figure 17 0. Micro­
cachrys tetragona
­foliage and seed cones

figure 172. Nageia nagi flushing leaves

f i g u re 1 7 1 . Nageia fleuryi leaves
and seed cones (photo L. Averyanov)

f i g u re 1 7 3. Nageia wallichiana tree in

­Vietnam (photo L. ­Averyanov)

figure 174. Nageia

wallichiana leaves and
seed cones (photo
L. Averyanov)

figure 17 5. Neo­
callitropsis pancheri figure 17 6. Neocallitropsis
in New ­Caledonia pancheri foliage
Juniperus sabina L. var. davurica (Pall.) Farjon, coast and in N Korea, but becomes increasingly
Monogr. Cupressaceae & Sciadopitys: 367. 2005. continental further west, with long and cold winters
Juniperus davurica Pall., Fl. Ross. 1 (2): 13, t. 55. and deep snow especially at higher altitudes. This
1789. Type: Russia: Russian Far East, Amur River, variety has also been found in a few localities in the
[“E Sibir. orientali”], P. S. Pallas [ex herb. Pallas] s.n. Alps, where conditions are similar to the localities
(lectotype BM). in mountains away from the coast in the Far East of
the species range.
Juniperus davurica Pall. subsp. maritima Urussov,
Bjull. Glavn. Bot. Sada 122: 55. 1981.
Conservation
IUCN: LC 457
Description
Decumbent shrubs to 50 cm. Branches ascending to
erect, whip shoots assurgent. Leaves of two types: ac- Juniperus saltillensis M. T. Hall, Fieldiania Bot.
icular and scale-like; acicular leaves usually present 34 (4): 45. 1971. Juniperus ashei J. T. Buchholz var.
from seedling to maturity, merging on some shoots saltillensis (M. T. Hall) Silba, Phytologia Mem. 7:
to scale leaves, in some populations scale leaves pre- 32. 1984. Type: Mexico: Coahuila, Sierra Madre
dominate. Pollen cones and seed cones invariably on Oriental, ca. 30 km SE of Saltillo, M. T. Hall 66305
branchlets with scale leaves. Dimensions of leaves (holotype F).
and cones not different from var. sabina.
Etymology
Taxonomic notes
The species epithet refers to Saltillo, a town in Coa-
The reduction to a variety of J. davurica Pall. under huila, Mexico, from the vicinity of which the species
the widespread species J. sabina L. is discussed and was first described.
explained in the Monograph of Cupressaceae and
Sciadopitys (Farjon, 2005a: 369).
Vernacular names
No common names are recorded for this species.
Distribution
China, Heilongjiang, Nei Mongol (Inner Mongolia);
Description
North Korea; Russia, E Siberia, Russian Far East;
Austria and probably occasionally elsewhere within Shrubs to 7 m, dioecious, sometimes monoecious;
the wide range of the species. trunk multistemmed, branching at ground level or
TDWG codes: 11 AUT-AU 30 (?) 31 AMU KHA PRM slightly above. Bark on larger stems relatively thick,
36 CHI-NM CHM-HJ 38 KOR-NK shredding, exfoliating in long, scaly or fibrous strips;
inner bark red-brown; outer bark grey. Branches nu-
merous, spreading in all directions, forming a bushy
Ecology
crown usually wider than tall. Foliage branches
Juniperus sabina var. davurica is a decumbent or low forming more or less dense tufts, rigid, (pen)ulti-
shrub in more or less open conifer forests with Lar- mate branchlets short, straight or curved distally,
ix gmelinii, Picea obovata or P. koraiensis. Outside 3–10(–14) mm long, 1.2–1.8 mm wide, ultimate
these forests it occurs along rocky streams as well as branchlets quadrangular in cross-section, older
on open slopes, sometimes between stands of Pinus branchlets more or less terete. Leaves decussate on
pumila, more often in alpine meadows and among ultimate branchlets, in alternating whorls of 3 on
rocks. Its main stem is usually buried in (peaty) top older branchlets and on whip shoots, imbricate, ap-
soil and in exposed situations it is extremely decum- pressed, all scale-like except on the most vigorous
bent. The altitudinal range is 400–1400 m a.s.l. Rock whip shoots, rhombic to ovoid-rhombic, 1.2–1.5 ×
types vary from granitic, metamorphic or volcanic 1–1.2 mm on ultimate branchlets, small leaves more
to calcareous; soils are usually peaty or leached and or less gibbous; margins minutely denticulate; apex
acidic. The climate is maritime temperate near the obtuse or acute; few stomata abaxially near base and
 scattered stomata adaxially from base to apex; glands Juniperus saltuaria Rehd. & E. H. Wilson, in
present on older leaves and most leaves of ultimate Sargent, Pl. Wilson. 2: 61. 1914. Type: China:
branchlets, not active, central or near leaf base, oval Sichuan, Min River, Songpan, mountains N of
and flat; leaf colour greyish green, sometimes yel- town, E. H. Wilson 3013 (lectotype K).
lowish green. Pollen cones terminal, solitary, ovoid-
oblong, 2–3.5 mm long; microsporophylls 8–10(–12),
Etymology
decussate, peltate; margins minutely denticulate,
keeled towards the obtuse apex, bearing 3–4 abax- The species epithet [Latin saltuarius = forester]
ial pollen sacs. Seed cones terminal on very short means ‘of mountain forests’.
branchlets, maturing in one year, when full grown
458 globose or ovoid-globose, 5–7 × 4.5–7(–8) mm,
Vernacular names
sometimes wider than long and slightly bilobed,
wine-red with thick whitish blue bloom (pruinose). fang zhi bai (Chinese)
Bract-scale complexes in 2 decussate pairs (total 4),
rarely including an upper whorl of 3 (total 5), com-
Description
pletely fused, with minutely exserted triangular
bract tips; surface smooth (rugose in sicco); fibrous Shrubs or trees to 15(–20) m, monoecious; multi-
and resinous inside. Seeds 1–2(–3) per cone, (broad) stemmed or monopodial; trunk to 1 m d.b.h. Bark on
ovoid or ovoid-oblong, 4–5(–6) × 3–3.5 mm, grooved larger stems exfoliating in longitudinal strips, weath-
or 2-ridged, with resin pits towards the base; hilum ering grey-brown or grey. Branches spreading, with
a third to half of length of seed, lighter than (dark) dense foliage forming a broadly rounded or irregu-
brown seed coat. lar crown. Foliage branchlets spreading, curved or
drooping, ultimate branchlets quadrangular, 1.0–1.7
mm wide and covered with appressed scale leaves,
Distribution
persistent. Leaves decussate, or 3-whorled on whip
Mexico: Chihuahua, Coahuila, Nuevo León, Zacate- shoots, imbricate, decurrent, on ultimate branchlets
cas. 1–2 × 0.7–1 mm, triangular-rhombic, gibbous, obtuse,
TDWG codes: 79 MXE-CO MXE-CU MXE-NL MXE- ridged abaxially, acute to pungent; weakly amphisto-
ZA matic, adaxial stomata in two bands separated by an
inconspicuous midrib; leaves on ultimate branch-
lets abaxially glandular; gland near the base, ovate
Ecology
to orbicular, convex or flat to slightly depressed; leaf
Juniperus saltillensis is usually an understorey shrub colour green, without cuticular wax. Pollen cones
in Pinyon-Juniper woodland or open Pinus-Quercus solitary, terminal on short branchlets, subglobose,
or Quercus forest, with Pinus cembroides, Juniperus 2–3 mm diam.; microsporophylls (4–)6–8, decussate,
spp., and Acacia sp., or in the ecotone to Bouteloua peltate-orbicular, with entire hyaline margins, bear-
grassland with e.g. Opuntia sp., Agave sp., and Ephe- ing 2–3 abaxial pollen sacs near lower margin. Seed
dra sp.; predominantly on limestone in the semi- cones terminal on short erect branchlets, maturing
arid zone (rainshadow) of mountain ranges. The in the second season to become globose or ovoid-
altitudinal range is from 1800 m to 2900 m a.s.l. ellipsoid, 5–8(–10) × 4–8 mm, lustrous blue-black
or purplish black and soft. Bract-scale complexes 6,
decussate, entirely fused; bract tips usually hidden or
Conservation
minute; surface smooth; scale tissue succulent, res-
IUCN: LC inous. Seeds 1 per cone, irregularly ovoid-globose,
3.5–7 × 3–5 mm, shallowly or deeply grooved, with
resin pits, yellowish to light brown.
Uses
No uses have been recorded of this species and it
Distribution
appears not to have been introduced to cultivation.
China: Gansu, Qinghai, Sichuan, NW Yunnan, E Xi-
zang [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHQ
Vernacular names
CHT
No common names are recorded for this species.
Ecology
Description
Juniperus saltuaria occurs in high ­montane to sub-
alpine coniferous forest, with Abies spp., Larix Small, shrubby trees to 5–8 m tall, often multi-
potaninii or Picea spp., and an understorey of stemmed, densely branched. Bark on branches
Rhododendron, Sorbus, and Salix; in more or less smooth, red-brown, soon flaking, thin, stringy, ex-
pure juniper groves or with J. convallium, J. pingii, foliation on stem(s) in strips or thin plates. Branches
J. ­squamata and Rhododendron spp. in the ecotone numerous, spreading, ascending or irregularly dis- 459
between forest and alpine meadows; and in the lat- posed, forming a dense, rounded or irregular crown.
ter in mixed shrub communities with e.g. Rhodo­ Foliage apparently in a fixed neotenic stage with only
dendron, Salix, Cotoneaster, Berberis, Lonicera, and juvenile leaves including fertile branchlets. Leaves
Spiraea. The altitudinal range of this species is from decussate, imbricate, decurrent, with proximal part
2100 m to 4600 m a.s.l. It is common in rocky areas, clasping shoot, distal part spreading 30–60° and
e.g. at the foot of limestone cliffs and crags and in much longer, total leaf length (4–)5–8(–9) mm, ca.
deep gorges, most abundantly on steep S-SW-facing 1 mm wide at the point of divergence, subulate-ac-
slopes, but occurs also in deeper mountain soils on icular; margins entire, slightly incurved; apex acute-
less steep slopes. It is not very drought tolerant and pungent; epistomatic, stomatal band undivided by a
needs abundant seasonal rainfall or snowmelt. weakly developed, proximal midrib; glands absent;
leaf colour lustrous mid-green with whitish stomatal
band. Pollen cones not observed. Seed cones termi-
Conservation
nal on very short ultimate branchlets, broadly ellip-
IUCN: LC soid when growing, becoming subglobose to slightly
reniform at maturity, 4–5 × 3–5 mm, dark blue with
white bloom (pruinose). Bract-scale complexes 4,
Uses
in two decussate pairs, not visible in mature cones.
This species is often too small to be a timber tree, but Seeds usually 2 per cone, sometimes slightly diver-
may be used for firewood or local carpentry in some gent apically, ca. 4 × 3 mm, ovoid-globose, light
parts of its natural range. It is rare in cultivation, re- brown.
stricted to a few arboreta or pineta, and not available
in the horticultural trade.
Taxonomic notes
Proposals to include J. saxicola as an infraspecific
Juniperus saxicola Britton & P. Wilson, Bull. Torrey taxon with J. barbadensis are apparently based on
Bot. Club 50: 35. 1923. Juniperus barbadensis L. sub- insufficient knowledge of the characters and traits of
sp. saxicola (Britton & P. Wilson) Borhidi, Acta Bot. these species. Adams (1995, 2000) studied the chem-
Acad. Sci. Hungarica 37 (1–4): 90. 1992; Juniperus istry of all relevant taxa in the Carribean and beyond,
barbadensis L. var. saxicola (Britton & P. Wilson) and also provided a key and descriptions to support
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 25. 2000. the distinctiveness of J. saxicola from J. barbaden-
Type: Cuba: Granma, Sierra Maestra, J. S. S. Léon sis. The most striking distinction is the retention of
10798 (holotype NY). juvenile type leaves with decurrent bases; this is a
feature rarely if ever found in any of the other sabi-
noid junipers of the western hemisphere. It is, on the
Etymology
other hand, not a reliable distinction at species level
The species epithet refers to its rocky habitat (Latin in sabinoid junipers of Eurasia (see e.g. J. chinensis,
saxum = rock, colere = to grow). J. sabina). Other characters that separate J. saxicola
 from its Carribean congeners are the distribution of
Etymology
stomata (epistomatic versus amphistomatic) and the
absence of leaf glands. Both may be traits of juvenile The species epithet means ‘of the rocks’, i.e. the spe-
leaves; so that the only real distinction may indeed cies grows in rocky places.
be the consistent formation of juvenile leaves and
the absence of scale leaves. The chemistry, whilst
Vernacular names
perhaps not giving conclusive evidence, at least in-
dicates that these plants, well separated geographi- Rocky Mountain juniper, Red-cedar, Rocky Moun-
cally from other junipers, are members of a separate tain red-cedar, River juniper; Cedro rojo (Spanish)
species.
460
Description
Distribution
Shrubs or small trees to 10–15(–20) m, dioecious,
Cuba (Granma, Sierra Maestra, Pico Turquino). rarely monoecious; multistemmed or monopodial,
TDWG codes: 81 CUB (short) trunk with a diam. up to 0.8–1(–2) m, rarely
a prostrate shrub in exposed rocky sites. Bark of
thick branches and trunk fibrous, rough, exfoliating
Ecology
in long, ragged strips or more compact, red-brown
This species is found at the higher altitudes in the weathering grey-brown. Branches ascending from
­Sierra Maestra, on ridges and rocky outcrops in a base or spreading, often contorted, those of higher
low (ca. 8 m), xeromorphic variant of the cloud for- orders ascending or spreading, forming a pyrami-
est, at altitudes between 1450 m and 1700 m a.s.l. dal or conical crown in young trees, to a broad,
Associated species are Cleyera ekmanii, Clusia tet- rounded or more irregular and open crown in old
rastigma, Haenianthus salicifolius, Lyonnia turquini, trees. Foliage branches numerous, irregularly dis-
and Ternstroemia microcalyx. The juniper occurs posed, often drooping to subpendulous in old trees,
where the rocky terrain prevents these angiosperms ultimate branchlets spreading to erect or droop-
from becoming dominant. ing, stiff or lax, slender, 5–35 mm long, 0.8–1.2 mm
thick, more or less quadrangular in cross section,
covered with scale leaves, persistent. Leaves decus-
Conservation
sate on ultimate branchlets, on whip shoots of young
Due to its very restricted areas of occurrence and trees sometimes ternate, imbricate, scale-like, 1–2 ×
occupancy this species, growing in a densely popu- 0.5–1 mm on lateral branchlets, decurrent, ovate-
lated region of Cuba, is considered to be at risk of rhombic; apex appressed, obtuse or mostly acute;
becoming extinct. margins entire; stomata on abaxial side limited to
IUCN: VU (D2) leaf base, on adaxial surface in two tapering bands;
glands conspicuous, large, elliptical or oblong,
slightly depressed, exudate often present; leaf colour
Uses
yellowish green to dark green. Pollen cones numer-
No uses have been recorded of this species; due to ous, terminal, solitary, ovoid or subglobose, 1.5–3.5
its shrubby habit it is unlikely to be exploited and × 1.5–2 mm; microsporophylls 6–8(–10), decussate,
firewood is available from other sources. It is not peltate, acute, with entire or erose upper margins
known to be in cultivation. and with 3–4 abaxial pollen sacs. Seed cones termi-
nal on straight or curved, short branchlets, maturing
to purplish ­orange or reddish brown with glaucous
Juniperus scopulorum Sarg., Silva N. Amer. 14: 93, or blue bloom in (1–)2 years, subglobose to trans-
t. 739. 1902. Type: USA: New Mexico, Santa Fe Co., versely broadened or bilobed, (5–)6–8(–9) mm wide
Santa Fe, A. Fendler 835 (lectotype K). (often wider than long), internally soft pulpy. Bract-
scale complexes 4–6, entirely fused, decussate,
Juniperus maritima R. P. Adams, Phytologia 89 (3): mostly indiscernible in mature cones, bract apices
278. 2007. minutely exserted, surface smooth, rugose in sicco.
Seeds 1–2(–4) per cone, with apices often diverging,
broadly ovoid or more or less flattened, 3.5–5 × 3.5–4 2300(–2770) m a.s.l. It is most common in rocky
mm, with rounded base and acutish apex, grooved terrain on soils derived from basalt, limestone and
and ridged on sides, pitted with resin concentrated shale, usually eroded and shallow, quickly drained
in the pits near the base, lustrous light yellowish and mostly nutrient-poor. The climate is mostly
brown, with a large darker hilum proximally. continental, dry (in the rain shadow of coastal
mountain ranges and in the SW) or subhumid with
scattered summer storms, and has extreme differ-
Taxonomic notes
ences in temperature between summer and winter.
Bilobed cones with two seeds are commonly found
in J. scopulorum (e.g. R. P. Adams 837 from North
Conservation
Dakota) and are not a character that is unique for 461
J. blancoi in North America. In Asia this character IUCN: LC
is most common and pronounced in J. semiglobosa,
but not rare in J. sabina and J. chinensis. Recently,
Uses
Adams (op. cit.) separated plants from islands in
Puget Sound, British Columbia and Washington, as This juniper is still being used as a source of fire-
a distinct species Juniperus maritima. The morpho- wood in rural areas especially by Native Americans;
logical data presented show no clear separation be- its use for fenceposts by ranchers has now almost
tween J. scopulorum and J. maritima and are in fact entirely been superseded by steel. It is in cultivation
nearly similar, with the exeption of some that may particularly in the USA, where a number of cultivars
have environmental causes. are known; several of these have reached Europe and
are there increasingly available in the trade. As with
other junipers, emphasis with these is on habit and
Distribution
less so on foliage colours. Due to its very wide latitu-
Canada: Alberta, British Columbia; USA: Arizona, dinal distribution certain provenances of the species
Colorado, Idaho, Montana, Nebraska, Nevada, New and at least some of its cultivars will be suitable for a
Mexico, North Dakota, Oregon, South Dakota, Tex- particular climate condition in temperate regions of
as, Utah, Washington, Wyoming; Mexico: Chihua- the world, ranging from the colds of Canada to the
hua, Coahuila. hot summers of Mexico.
TDWG codes: 71 ABT BRC 73 COL IDA MNT ORE
WAS WYO 74 NEB NDA SDA 76 ARI NEV UTA 77
NWM 79 MXE-CO MXE-CU Juniperus semiglobosa Regel, Trudy Imp. S.-Peter-
burgsk. Bot. Sada 6 (2): 487. 1879. Type: Tadjikistan:
Zaravshan Range, Saratag Pass, Isfara, B. A. Fedt-
Ecology
schenko s.n. (lectotype LE). Fig. 147, 148, 149
A major component of the Rocky Mountain foot-
hills in a woodland coniferous zone (in the south- Juniperus media V. D. Dmitriev, Trudy Sektora Ag-
ern portions of its range Pinyon-Juniper woodland, rolesomelior. Lesn. Khoz. Komiteta Nauk Uzbeksk.
with Pinus edulis and Juniperus osteosperma) and SSR 4: 31. 1938.
extending upwards into montane coniferous forest Juniperus tianschanica Sumnev., Bot. Mater. Gerb.
(mainly with Pinus contorta, P. ponderosa, P. flexilis, Inst. Bot. Zool. Akad. Nauk Uzbeksk. SSR 8: 24.
in the north also Picea engelmannii, Abies lasiocar- 1946.
pa), where it often occupies exposed rocky slopes
and ridges. Downslope it extends into scrubland
Etymology
often dominated by Seriphidium tridentatum (Ar-
temisia tridentata), with e.g. Amelanchier spp., Chry- The species epithet describes the ‘half-round’ shape
sothamnus spp., and Prunus spp., and into Quercus of the seed cones.
woodland; along streams it occurs on dry sites in
Populus-Salix woodland. Only in the northern parts
Vernacular names
of its range, at lower and middle elevations, does it
form pure stands. The altitudinal range is (5–)500– Pencil cedar; kun lun duo zi bai (Chinese)
 Description Ecology
Arborescent shrubs or trees to 15(–20) m tall, rarely In (mixed) conifer forests with Abies pindrow,
decumbent, dioecious or rarely monoecious; trunk ­Cedrus deodara, Picea schrenkiana or Pinus wal-
to 1.2 (–2) m diam. monopodial or with several lichiana, usually on open rocky slopes or outcrops;
stems low above ground. Bark on large trunks fis- above this zone and on S-facing slopes extending
sured, fibrous, exfoliating in long strips, brown into a juniper woodland type (often scattered) with
weathering grey-brown. Branches long, spreading J. excelsa subsp. polycarpos or J. pseudosabina, and
and ascending, in young trees nearly erect; crown on glacial moraines, where it can form groves. It also
variable from pyramidal to broad and irregular in occurs in subalpine scrubland with J. pseudosabina,
462 old trees. Foliage branchlets numerous, slender to J. sabina, J. communis var. saxatilis, Fraxinus sp.,
thick, terete or sometimes slightly quadrangular, ul- Salix spp., Origanum, Pteridium, Rosa, Saussurea,
timate branchlets to 4 cm long, 1–2.5 mm thick, cov- Scabiosa, etc., extending into interior valleys and
ered with scale leaves, persistent. Leaves decussate semi-arid high plains dominated by Seriphidium
on ultimate branchlets, closely appressed, scale-like, maritimum (Artemisia maritima). The altitudinal
1–2 × 1–1.2 mm, ovoid-rhombic to nearly triangu- range is 1550–4350 m a.s.l. It occurs on river terraces,
lar, often ternate; margins hyaline, entire; abaxially moraines, scree slopes and other dry, stony ground,
a few stomata near base and in two tapering bands both calcareous and siliceous. The climate is alpine-
adaxially; glands central, large and conspicuous, el- continental, with hot, dry summers especially in the
liptic, often active; leaf colour lustrous light or yel- interior valleys and on S-facing mountain slopes,
lowish green with whitish cuticular wax, sometimes and cold winters, often with perpetual snowfields
glaucous green. Pollen cones numerous, terminal above the juniper zone supplying moisture through
on ultimate branchlets, subglobose, 3–5 × 2–4 mm; prolonged melting and seepage.
microsporophylls 8–10, decussate, peltate, with en-
tire hyaline margins, bearing 3–4 abaxial pollen sacs
Conservation
near lower margin. Seed cones terminal on branch-
lets, growing in two seasons from green or purplish IUCN: LC
to light brown to dark blue with a glaucous white
bloom, caducous, subglobose to (tri)angular, 4–8
Uses
× 4–6 mm, pulpy or fibrous, more or less resinous,
maturing dry and hard, sometimes more succulent. This species is primarily used for firewood, but the
Bract-scale complexes 4(–6), usually decussate or wood is used on a small scale for wood craft (sou-
sometimes (in part) in whorls of 3, entirely fused, venirs) and the foliage is sold on markets for ‘me-
smooth, rugose when dry, with triangular bract api- dicinal’ purposes (pers. obs. in Kirgyzstan, August
ces protruding 0.3–0.5 mm. Seeds (1–)2–3(–4) per 2000). In horticulture Pencil cedar is uncommon
cone, 3–6 × 2–3.5 mm, angular, subovoid or conical, and it may on occasion have been introduced as
yellowish brown or red-brown, with a large proxi- J. polycarpos or J. macropoda, depending on what
mal hilum. part of the range of J. semiglobosa such mis-identi-
fications originated from. It is probably confined to
collections in arboreta and pineta if at all present in
Distribution
gardens; very few plants are recorded and may not
Central Asia: Kazakhstan (near Chimkent), Kir- have survived where they were grown.
gyzstan, Tadzhikistan, Uzbekistan; W Asia: Af-
ghanistan, Pakistan (Baltistan, Gilgit Wazarat,
Hazara, Hindu Kush, Karakoram Range, Kohistan); Juniperus squamata Buch.-Ham. ex D. Don, in
China: Xinjiang, W Xizang [Tibet]; India: Himachal Lambert, Descr. Pinus 2: 17. 1824. Type: Bhutan:
Pradesh, Jammu-Kashmir; Nepal. [locality unknown (“Habitat in Bhotaniae
TDWG codes: 32 KAZ KGZ TZK UZB 34 AFG 36 Alpibus”)], W. S. Webb W 6043C (lectotype K-W).
CHT CHX 40 NEP PAK WHM-HP WHM-JK
Juniperus morrisonicola Hayata, Gard. Chron., ser.
3, 43: 194. 1908; Juniperus squamata Buch.-Ham.
ex D. Don var. morrisonicola (Hayata) H. L. Li & covered with whorls of small scale leaves, growing in
H. Keng, Taiwania 5: 81, t. 28. 1954. two seasons via a green phase to bluish black, sub-
Juniperus squamata Buch.-Ham. ex D. Don var. globose to ovoid, 4–8 × 4–6 mm, soft and succulent.
fargesii Rehd. & E. H. Wilson, in Sargent, Pl. Wilson. Bract-scale complexes 3, of equal size, fused with in-
2: 59. 1914. visible margins; outer surface smooth, lustrous, with
Juniperus baimashanensis Y. F. Yu & L. K. Fu, Novon 3 subapical bract tips just visible or hidden. Seeds
7 (4): 443. 1998. ovoid-globose, light brown, 3.5–6 × 2–5 mm with
Juniperus squamata Buch.-Ham. ex D. Don var. several faint, darker resinous grooves or pits.
parvifolia Y. F. Yu & L. K. Fu, Novon 7 (4): 444. 1998.
Juniperus squamata Buch.-Ham. ex D. Don var.
Distribution
hongxiensis Y. F. Yu & L. K. Fu, Novon 7 (4): 444. 463
1998. Afghanistan; N Pakistan; India (Himalaya); Ne-
pal; Bhutan; N Myanmar [Burma]; China: S An-
hui, Chongqing, W Fujian, S Gansu, E Guizhou, W
Etymology
Hubei, Shaanxi, Sichuan, Xizang [Tibet], Yunnan;
The species epithet means ‘scaly’ or ‘with scales’ and Taiwan.
presumably refers to the flaky bark. TDWG codes: 34 AFG 36 CHC-CQ CHC-GZ CHC-
HU CHC-SC CHC-YN CHN-GS CHN-SA CHQ CHS-AH
CHS-FJ CHT 38 TAI 40 ASS-AS EHM-AP EHM-BH
Vernacular names
EHM-DJ EHM-SI NEP PAK WHM-HP WHM-JK WHM-
Nepalese juniper, Himalayan juniper, Flaky juniper; UT 41 MYA
Pudma Chundur (India); gao shan bai (Chinese)
Ecology
Description
Occuring from subalpine coniferous forest and
Shrubs, procumbent at high altitudes, or small trees mixed woodland with Abies spp., Picea spp., Larix
10–12 m tall, monoecious; rarely monopodial, usu- spp. and Juniperus semiglobosa, J. recurva, J. saltuar-
ally multistemmed or branching near the ground. ia, Betula spp., and Quercus spp., up to subalpine
Bark on larger stems becoming fissured, exfoliating Rhododendron thickets and Juniperus thickets or
in long, thin strips, turning grey-brown. Branches alpine dwarf shrub or grass/forb communities. In
thick and short esp. in mountain forms, twisted, thickets and alpine scrub it is commonly associated
numerous in alpine shrub forms, spreading or as- with Juniperus indica, J. pingii var. wilsonii, Berberis,
cending. Foliage branches densely crowded or more Caragana (in NE of range), Cotoneaster, Polygonum
sparse, patent, ultimate branchlets initially triangu- bistorta, Rhododendron, Rosa, Sorbus, Spiraea etc.
lar in cross section, green, becoming more or less The altitudinal range is 1340–4850 m a.s.l. It is found
terete, reddish brown, persistent. Leaves in alternate on various rock types, from calcareous to siliceous,
whorls of 3, decurrent, free part (distal half to two and often predominant on moraines, scree slopes or
third of total length) more or less spreading to pat- rocky ridges, but also on gravelly flood plains. The
ent, acicular, subulate-lanceolate to linear, straight climate is high montane to alpine with strong mon-
or slightly curved, (2.5)4–10 × 0.8–1.5 mm, widest soon influence, which however diminishes towards
below middle part; margins entire; apex acute or the NE of its range.
acuminate; epistomatic, stomata in two bands sepa-
rated by a narrow midrib often concealed by a cov-
Conservation
ering of white cuticular wax; abaxial surface with a
thin groove or ridge; leaf colour green to glaucous IUCN: LC
green. Pollen cones terminal on axillary dwarfed
shoots, solitary, subglobose to ovoid, 2–2.5 × 3–4
Uses
mm; microsporophylls 9–12, ternate, more or less
cordate-peltate, obtuse or cuspidate, with entire, Juniperus squamata is widely cultivated as a garden
hyaline margins, abaxially bearing 3 ovoid pollen ornamental and several shrubby and prostrate forms
sacs. Seed cones terminal on axillary dwarfed shoots are propagated as cultivars. Forms with ­glaucous
 leaves are much in demand and accordingly a con- epistomatic, with two stomatal bands separated by
stant stream of new cultivars with this trait runs a narrow midrib. Leaf colour light green or glau-
into the horticultural market. Juniperus squamata cous green, leaves at tips of branchlets yellowish
‘Wilsonii’ is not a cultivar of this species, but the green. Pollen cones numerous, terminal, on usually
taxon J. pingii var. wilsonii. recurved branchlets, solitary, subglobose or ovoid,
2–3 × 1.5–1.8 mm; microsporophylls 6–10, decussate,
peltate, more or less gibbous, with rounded, erose-
Juniperus standleyi Steyerm., Publ. Field Mus. Nat. hyaline margins, abaxially bearing 3–4 pollen sacs.
Hist., Bot. Ser. 23 (1): 3. 1943. Type: Guatemala: San Seed cones terminal on very short, curved branch-
Marcos, Volcán de Tacaná, J. A. Steyermark 36137 lets, growing in one year to subglobose (5–)6–7(–9)
464 (holotype F). mm diam., purplish brown or brown, often with a
glaucous bloom. Bract-scale complexes 6, two up-
per pairs entirely fused, surface smooth or rugose
Etymology
(in sicco), with minute bract apices of upper scales
This species was named after the botanist P. C. Stand­ protruding; cone tissue dry, dense, resinous. Seeds
ley, co-author (with J. A. Steyermark) of the Flora of (2–)3–4 per cone, broadly ovoid, 2.3–3.5 × 2–2.5 mm,
Guatemala. more or less angular, with grooves and pits filled
with hard resin, brown with lighter hilum.
Vernacular names
Distribution
Huitó, Huitum; Cipres (Spanish)
Mexico: Chiapas (Volcán Tacana); Guatemala: Hue-
huetenango, San Marcos (highlands).
Description
TDWG codes: 79 MXT-CI 80 GUA
Decumbent to erect dioecious shrubs or trees of me-
dium to large size, trees to 15–20 m, multi-stemmed
Ecology
or monopodial. Bark becoming rough and scaly,
exfoliating in long flakes, at first reddish brown or This species occurs in open pine woodland (Pinus
purplish brown, turning dark brown. Branches nu- spp.), or sometimes in pure stands, often on rocky
merous, spreading or ascending, foliage branches edges and talus slopes of mesas, or on limestone
mostly ascending, on decumbent plants creeping ridges. It grows there with shrubs, grasses, and
over rocks, forming a spreading or bushy, dense forbs, sometimes on land under some extensive cul-
crown. Foliage branchlets mostly 5–15 mm long, tivation (Agave). The altitudinal range is from 3000
sometimes longer to 20 mm, 1.1–1.4 mm wide, m to 4250 m a.s.l. At its upper limit it occurs in high
quadrangular in cross-section, straight or curved. altitude pine forest with Pinus hartwegii, where in
Adult type leaves (scale leaves) decussate, on (pen) some places it reaches the tree line.
ultimate branchlets appressed and imbricate, with
smallest leaves 1–1.5(–1.7) × 0.7–1.1 mm; on older
Conservation
branches leaves substantially larger and becoming
acute, with free or recurved apex, acute-mucronate, This species is geographically restricted to an area of
on whip shoots ovoid-rhombic, gibbous; margins ca. 700 km2, but within that range less than a third
entire, incurved; apex obtuse or rounded. Stomata of the original stock historically present remains
mostly adaxially, a few abaxially on two sides near (Islebe, 1993). The scarcity has led to municipal
margins of leaves; glands inconspicuous, in a slight boundary disputes. Official permission is now need-
depression below leaf centre, usually inactive, but ed to cut the trees (shrubs are not used) but the local
on whip shoots often producing a drop of resin, inhabitants largely ignore this rule. In the rainy sea-
becoming oblong in elongated leaf shape. Juvenile son there is extensive sheep grazing on the altiplano,
type leaves recurring on mature trees after dam- which prevents successful regeneration.
age and resprouting, lanceolate-subulate, navicu- IUCN: EN (B1+2b)
late, stiff, (4–)5–7 × 1 mm (at base), acute-pungent;
 and thick, adaxial midrib mostly continuous to the
Uses
obtuse, or occasionally acute apex; epistomatic, sto-
The local Todosanteros Indians use the wood for mata in two conspicuous bands separated by a thin,
fence posts and shingles. Smaller wood may serve as raised midrib; leaf colour light green or glaucous,
firewood. This rare species does not appear to be in with white stomatal bands. Pollen cones axillary,
cultivation; its precarious conservation status mer- solitary, ovoid to cylindrical, obtuse, ca. 5 × 3 mm;
its ex situ conservation efforts, which include taking microsporophylls 9–12, in alternating whorls of 3,
the species into cultivation. Due to its high altitude imbricate, subpeltate, broadly triangular, weakly
habitat it should prove hardy in temperate regions keeled; margins entire, bearing 5–6(–7) abaxial pol-
of the world. len sacs. Seed cones axillary on very small dwarf
shoots with 2–3 whorls of small scale leaves, matur- 465
ing in 2 years, becoming subglobose, often more or
Juniperus taxifolia Hook. & Arn., Bot. Beechey less triangular towards apex, 8–10 mm diam., chang-
Voy. 6: 271. 1838. Type: Japan: Bonin Islands, ing from orange-brown to reddish brown or more
Ogasawara Group, [“Bonin”], G. T. Lay & A. Collie often purplish red or dark purplish blue with a light
s.n. (holotype K). blue bloom (pruinose). Bract-scale complexes in 2–3
alternating whorls of 3, upper whorl much enlarged,
Juniperus lutchuensis Koidz., Bot. Mag. (Tokyo) 32: completely fused but sutures visible near distal pole
138. 1918. of the cone; upper 3 bract tips exserted, minute; cone
tissue soft pulpy, succulent when ripe. Seeds 3 per
cone, ovoid-conical, 4.5–6 × 3 mm, more or less tri-
Etymology
angular in cross-section, oblique; resin grooves and
The species epithet means ‘with leaves as of Taxus’ pits conspicuous; apex mucronate; seed colour light
(yew). brown or tan.

Vernacular names Distribution

Luchu juniper, Yew-leaved juniper; Hainezu, Japan: Ryukyu Islands (Okinawa Group, Amami
Okinawa-hainezu, Fitcheisi (Japanese) Group); NW Pacific (admin. by Japan): Ogasawara
Group [Bonin Is.]. Also reported from a locality in
Honshu, but it is uncertain if it is indigenous there.
Description
TDWG codes: 38 NNS OGA
Decumbent to erect shrubs or small trees to 3–4 m,
dioecious; trunk or stems prostrate, or erect and
Ecology
multistemmed, rarely monopodial, up to 20 cm
diam. Bark on stems with papery flakes, grey-brown. Juniperus taxifolia forms a prostrate or decumbent
Branches spreading horizontally, rooting in decum- shrub close to the seashore above the tide mark, on
bent forms, often covering large areas, or assurgent, rocks or rocky slopes from sea level to the highest
forming a variable crown from decumbent to erect parts of the islands. It is also present further inland,
or more or less conical. Foliage branches spreading where it can be a small, erect tree, in tall grassland
or upright on decumbent plants, drooping or sub- and thickets on deeper, though usually rocky or
pendulous on (older) shrubs to small trees, young gravely soils. It is obviously subjected to strong and
shoots triangular in cross-section, becoming terete moist, salt-laden ocean winds.
with age, persistent, internodes 1–3 mm long, on
erect plants with drooping foliage to 6 mm. Leaves
Conservation
in alternating whorls of 3, non-decurrent, sessile, ar-
ticulate at base, spreading forward or at right angle In the Status Survey and Conservation Action Plan
from shoot, acicular-linear, 7–12(–16) × 1–1.5(–1.8) for Conifers of IUCN/SSC (Farjon & Page, 1999),
mm, straight or more often curved at base, with par- this species is listed with other conifers that had
allel sides or slightly wider towards base, obtusely been earlier listed as endangered (Farjon et al.,
keeled abaxially, concave adaxially; margins entire 1993). Juniperus taxifolia occurs on isolated oceanic
 islands and possibly in one locality on the beach or more dense, forming a pyramidal crown, in old
of a pen­insula in Honshu. Its extent of occurrence shrubs and trees more rounded and wider than tall.
(EOO) and area of occupancy (AOO), if calculated Foliage branches spreading or drooping to subpen-
separately for each island (this was not done) could dulous, branchlets slender or stout, 1–1.3 mm thick,
fall within the threshold for EN, while it is known more or less quadrangular in cross-section, covered
from fewer than 10 localities if each island is taken with scale leaves, persistent. Leaves decussate, im-
as a locality, but there is no evidence of threats and/ bricate, scale leaves on ultimate branchlets all ap-
or decline. pressed, on older branchlets some with spreading
IUCN: LC apex, on ultimate branchlets 1.3–2.7 × 0.7–1 mm,
ovate-rhombic, more or less gibbous or keeled dis-
466 tally; margins entire or hyaline-erose; apex obtuse
Uses
or acute; stomata on abaxial side in two small areas
In some of the larger islands (e.g. Okinawa) this near base, on adaxial side in two bands converging
species is being used as a shore windbreak and also to apex, covered with white or glaucous wax; gland
cultivated in gardens. It is in cultivation in gardens conspicuous or inconspicuous, mostly inactive,
in Japan, but rare. In the past its wood was used to central on rhombic scale leaves, in slight depres-
a limited extent in house building, for posts and as sion, oval to oblong; leaf colour light green. Pollen
fuel. Seeds were sent to the Arnold Arboretum in cones terminal, solitary, subglobose, 2–3 mm long;
Massachussetts, USA by Ernest Wilson in the early microsporophylls 10–12, decussate, peltate, con-
part of the 20th century; it remains a rarity in hor- vex, with rounded, hyaline-erose margins, bearing
ticulture outside the islands where it is also native. (2–)3–4 abaxial, angular-subglobose pollen sacs.
Seed cones terminal on straight or slightly curved
ultimate branchlets, growing in two seasons becom-
Juniperus thurifera L., Sp. Pl. 2: 1039. 1753. ing globose or subglobose, 7–8(–10) mm diam., fi-
Type: Spain: Teruel, Mansana, Camarena River, nally blackish blue or purplish black. Bract-scale
E. Reverchon 788 (neotype BM). complexes 4–6, decussate, completely fused; surface
usually smooth; bract tips exserted, less than 0.5
Juniperus thurifera L. var. africana Maire, Bull. Soc. mm, with sutures of individual scales often visible;
Hist. Nat. Afrique N. 17: 125. 1926; Juniperus africana scale tissue hard and dry pulpy. Seeds (1–)2–3 per
(Maire) Villar, Types Sols Afrique N. 1: 91. 1947; Ju- cone, irregular in size and shape in a single cone,
niperus thurifera L. subsp. africana (Maire) Romo & 3–5 × 3–4.5 mm, triangular or flattened, with shal-
Borantinsky, Ann. Bot. Fennici 44: 73. 2007. low pits and grooves, lustrous light brown, with a
dull tan hilum at base.
Etymology
Distribution
The epithet is from Latin turifer = yielding or pro-
ducing incense. SW Europe: S France (Alpes Maritimes), N and SE
Spain; North Africa: Morocco, N Algeria.
TDWG codes: 12 FRA-FR SPA-SP 20 ALG MOR-MO
Vernacular names
Spanish juniper, Incense juniper; Cedro de España,
Ecology
Trabina, Sabina blanca, Enebro (Spanish); Aoual,
Taoualt, Araâr (Algeria, Morocco) In semi-arid woodland with Quercus ilex; on the
Saharan slopes of the Haut Atlas in Morocco with
Cedrus atlantica and Quercus ilex, but extending
Description
above these in groves with only very sparse scrub
Shrubs or trees to 10–15(–20) m tall, dioecious; vegetation. The altitudinal range is (300–)900–
trunk monopodial or branching near the ground, to 2500(–3300) m a.s.l. At middle altitudes (1800–2500
1.5–2 m. d.b.h. Bark on trunks scaly, exfoliating in m) with e.g. Adenocarpus anagyrifolius, Ephedra
strips, dark brown, weathering grey-brown. Branch- major, and Fraxinus xanthoxyloides, at highest alti-
es spreading or ascending, numerous, foliage open tudes (to 3300 m) with Alussum spinosum, Prunus
prostrata, Bupleurum spinosum, Berberis hispanica,
Description
and Daphne laureola. In Spain it has a prevalence for
calcareous soils, e.g. on marls, but in the Atlas Mts. Trees or shrubs to 15 m, usually 5–10 m, monoecious,
it seems indifferent to soil type, usually growing on rarely dioecious; multistemmed or monopodial;
very rocky, skeletal soils. The climate is continental, trunk up to 2 m d.b.h. Bark on larger stems fissured
semi-arid, with cold winters (Atlas Mts.) to conti- and exfoliating in longitudinal fibrous strips, cinna-
nental-Mediterranean, with long dry summers. mon-brown, weathering grey. Branches ascending
or spreading, foliage branches very dense, short,
stiff and spreading at highest altitudes, elsewhere
Conservation
drooping or nearly pendulous, forming a dense,
This species is not considered to be threatened with broad pyramidal to eventually rounded or irregular 467
extinction. However, in Europe gradual alterations crown. Foliage branchlets ultimately stout, subterete,
of woodlands due to resumed forest succession after 1–1.5(–2) mm wide, 0.5–2 cm long, covered with ap-
abandonment of traditional land use systems could pressed leaves, persistent. Leaves on mature plants
eventually reduce the abundance of the juniper com- scale-like, decussate or in alternate whorls of 3, im-
ponent. Conversely, in the Haut Atlas of Morocco bricate, decurrent, 1.5–3 × 1–1.5 mm, ovate-rhombic,
heavy grazing and browsing pressures have caused obtuse; margins entire; stomata in 2 lines on each
damage and prevent regeneration. side mostly near base; gland abaxial and central in a
IUCN: NT depression or groove, elliptic or oblong, sometimes
inconspicuous, weakly active; leaf colour green or
yellowish green. Pollen cones numerous, solitary,
Uses
terminal on short branchlets, globose to subglo-
This species has traditionally been utilised in France bose, 1.5–2 mm diam.; microsporophylls 6–8, dec-
and Spain for construction wood, firewood, and spe- ussate, peltate-cordate, with rounded entire hyaline
cial uses (fences, various farming tools). The foliage margins, bearing 2–3 abaxial pollen sacs near lower
served as fodder for donkeys and goats and was, and margin. Seed cones terminal on short branchlets,
perhaps still is, used in Spain for incense. The use maturing in the second season to become globose
as animal fodder is still prevalent in North Africa. or ovoid, 10–16(–18) × 7–13(–14) mm, lustrous red-
In horticulture Spanish juniper is virtually limited dish brown or brownish black to purplish black, soft.
to botanic gardens and arboreta, mainly in south- Bract-scale complexes 4–6, decussate, entirely fused;
ern Europe, but some specimens are grown or have bract tip subapical, triangular, 1–1.5 mm; surface
been grown as far north as Ireland and the south of smooth; scale tissue succulent, resinous. Seeds 1 per
England. cone, subglobose to ovoid, or acutish at both ends,
laterally compressed, 7–11 × (5–)6–8 mm, ridged and
grooved, with deep resin pits, yellowish brown.
Juniperus tibetica Kom., Bot. Mater. Gerb.
Glavn. Bot. Sada RSFSR 5: 27. 1924. Type: China:
Distribution
Xizang (Tibet), Jinsha River, in forest near temple
[“Tschunkor gomba”], V. F. Ladygin 25 (holotype China: S Gansu, SE Qinghai, Sichuan, S and E Xi-
LE). zang [Tibet].
TDWG codes: 36 CHC-SC CHN-GS CHQ CHT
Juniperus distans Florin, Acta Horti Gothob. 3: 6.
1927.
Ecology
Juniperus tibetica is a species that forms groves or
Etymology
small forests at high altitudes, often together with
The species epithet denotes its origin in Tibet. J. convallium. The altitudinal range is 2600–4780
(–4900) m a.s.l. These woods are often grazed by
cattle (yaks) and degraded, being transformed into
Vernacular names
parkland with mats of Cyperaceae and low grasses
Tibetan juniper; da guo yuan bai (Chinese) and herbs, or further to Berberis-Juniperus scrub and
 Artemisia steppe. At lower altitudes it may occur in Vernacular names
clearings of Picea forest, usually on S-facing slopes; at
Eastern Redcedar, Pencil cedar, Red juniper, Red­
the tree limit, where it may be the only species, it re-
cedar; Savin, Cèdre rouge (French)
turns to S-facing slopes indicating that temperature
is the limiting factor. Locally it occurs in open stands
Description
of Cupressus torulosa var. gigantea. It occurs on rocky
slopes and ridges, as well as old gravel terraces, on Trees or arborescent shrubs, usually 15–20 m, ex-
rocky soils from siliceous as well as calcareous parent ceptionally to 25–27 m tall, dioecious; monopodial
material. The climate, mostly in the monsoon shad- or multistemmed; trunk with a maximal diam. of
ow of the main crest of the Himalayas, is continental 1–1.2(–1.7) m, often fluted at base and forked in
468 with extremes of solar radiation and frost. crown. Bark of trunk very fibrous, shredding, exfo-
liating in long, ragged strips, light red-brown weath-
ering grey-brown. Branches ascending or spreading
Conservation
in older trees, forming a pyramidal or conical crown
This species is widespread and not threatened with usually in young trees but locally in mature trees, to
extinction in the wild. There is, however, consider- a broad, rounded or more irregular and open crown
able pressure from both direct uses by indigenous in old trees. Foliage branches numerous, ascending
peoples (firewood in particular) as from grazing or spreading, often drooping to subpendulous in old
by domestic animals, which are likely to increase trees, ultimate branchlets spreading or drooping,
and have a detrimental effect on regeneration and lax, slender, 5–25 mm long, 0.8–1 mm thick, quad-
possibly its long term survival in many areas (G. & rangular or subterete in cross section, covered with
S. Miehe, pers. comm.). scale leaves, persistent. Leaves decussate on ultimate
IUCN: NT branchlets, scale-like, 1.3–3 × 0.7–1 mm, decurrent,
ovate-rhombic; apex usually free, acute or apiculate;
margins entire; stomata on abaxial side limited to
Uses
leaf base, on adaxial surface in two tapering bands;
As the principal high altitude tree in large parts of glands conspicuous, mostly well below middle of
Xizang [Tibet] and adjacent areas, this species is visible part of leaf, large, elliptical to circular, exu-
an important source of wood to local people, who date often present; leaf colour light green to dark
use it for firewood and on a limited scale for other green or glaucous green. Pollen cones numerous,
purposes, such as incense in Buddhist rituals. In- terminal, solitary, ovoid or oval, 2–3 × 1.5 mm; mi-
creased population pressures in recent decades have crosporophylls (8–)10–12, decussate, peltate, acute,
resulted in shortages of a resource that renews it- with entire or erose upper margins and with 3–4
self only very slowly. It is rare in cultivation; seeds abaxial pollen sacs. Seed cones terminal on straight,
were collected by Joseph Rock in southern Gansu in short branchlets, maturing to purplish blue with
1926 (and distributed under the synonym J. distans); glaucous bloom in (1–)2 years, subglobose to ovoid,
plants raised in the UK grew into a columnar habit 4–6.5 × 3–5.5 mm, internally soft pulpy, aromatic.
with light green leaves. Recent seed collections from Bract-scale complexes 4–6, entirely fused, decus-
Tibet by Georg and Sabine Miehe have germinated sate, mostly indiscernible in mature cones, bract
in the Botanic Garden of the University of Göttin- apices minutely exserted, surface smooth, rugose
gen in Germany. This tree forming species should be when dry, usually strongly glaucous. Seeds 1–2(–4)
grown more widely. per cone, broadly ovoid-conical or more or less flat-
tened on one side, 3.5–5 × 3.5–4 mm, with broad
base and acutish apex, grooved and ridged on sides,
Juniperus virginiana L., Sp. Pl. 2: 1039. 1753. pitted with resin concentrated in the pits near the
base, light yellowish brown, with a darker apex.
Etymology
Taxonomic notes
The species epithet means ‘from Virginia’; in the
time of Linnaeus used for much of what is now the In the Checklist of United States Trees (Little, 1979)
SE USA. and in Silvics of North America (Burns & Honkala,
1990) Juniperus silicicola (Southern Redcedar) and if the distinct leaf arrangement holds true in culti-
J. virginiana (Eastern Redcedar) were treated as dis- vars, juvenile leaves on lateral branchlets beyond the
tinct species. In Flora of North America 2 (1993), seedling stage of J. virginiana should be in pairs, not
Adams treated the former as a variety of the latter. in threes. Scale leaves of J. virginiana are acute, of
In Flora of Florida (Wunderlin & Hansen, 2000), J. chinensis usually obtuse.
J. silicicola has been sunk into the synonymy of
J. virginiana. Ecologically, there appears to be a dis- 2 varieties are recognized:
tinction which has been observed in other species in
the genus as well: a form largely restricted to coastal Juniperus virginiana L. var. virginiana. Type: USA:
sand dunes and sand flats as opposed to popula- locality unknown, leg. ign. LINN 1198.7 (lectotype
tions further inland on different soil types. Numer- LINN). Fig. 150 469
ous reports on hybridization and introgression with
J. horizontalis, which borders on J. virginiana in the
Description
north, and with J. scopulorum in the west are gener-
ally confirmed by subsequent research, but hybridi- Tree habit variable, including fastigiate branching
zation with J. ashei in the southwest has been refuted and conical crown but often broad and rounded
in subsequent studies (Adams, 1986). in mature trees. Leaves 1.5–3 × 0.8–1.2 mm, acute-­
apiculate or apiculate on whip shoots.
Distribution
Distribution
E North America: from S Quebec to Florida, west-
ward to North Dakota and Texas. Canada: SE Ontario, S Quebec, USA: Arkansas,
TDWG codes: 72 ONT QUE 74 ILL IOW KAN MIN Connecticut, Delaware, D. C., N Florida, Georgia,
MSO NDA NEB OKL SDA WIS 75 CNT INI MAI MAS Illinois, Indiana, Iowa, Kansas, Kentucky, Louisi-
MIC NWH NWJ NWY OHI PEN RHO VER WVA 77 TEX ana, Maine, Maryland, Massachussetts, Michigan,
78 ALA ARK DEL FLA GEO KTY LOU MRY MSI NCA Minnesota, Mississippi, Missouri, Nebraska, New
SCA TEN VRG WDC Hampshire, New Jersey, New York, North Caroli-
na, North Dakota, Ohio, Oklahoma, Pennsylvania,
Rhode Island, South Dakota, Tennessee, Texas, Ver-
Ecology
mont, Virginia, West Virginia, Wisconsin.
This species occurs in a wide range of habitats, from TDWG codes: 72 ONT QUE 74 ILL IOW KAN MIN
old, eroded sandstone or limestone plateaux covered MSO NDA NEB OKL SDA WIS 75 CNT INI MAI MAS
in open pine or pine-oak woodland, or stream banks MIC NWH NWJ NWY OHI PEN RHO VER WVA 77 TEX
of clay or sand in the Midwest, to abandoned fields 78 ALA ARK DEL FLA GEO KTY LOU MRY MSI NCA
and road verges and stabilised sand dunes on the At- SCA TEN VRG WDC
lantic coast. The altitudinal range is 1–1000 m a.s.l.
More details are given under the two varieties.
Ecology
Juniperus virginiana var. virginiana is a very wide-
Uses
spread conifer in the eastern United States, extend-
This species is used as an amenity tree in landscap- ing both its area of extent (total range) and its area
ing and as an ornamental. The wood is also exploited of occupancy. This happens primarily through colo-
on a small scale for special purposes such as wood nisation on disturbed ground which is subsequently
turning. In the past it was exploited extensively for left to settle; foremost are abandoned arable fields
the manufacture of pencils, but more recently that and pastures, second in importance are verges of
use has declined. From the leaves essential oils are highways, old mine tailings, etc. Other tree species
extracted. Juniperus virginiana has been the source invading these fields are Pinus spp., Ulmus ameri-
of numerous cultivars, many of which are still in the cana, Populus grandidentata, Sassafras albidum, and
horticultural trade. It is similar in general appear- Zanthophyllum americanum. In the primary range of
ance to J. chinensis and if they do not bear cones, e.g. this more continental and upland variety, the most
because of the retaining of juvenile leaf types, and commonly associated trees are Pinus ­virginiana,
 P. echinata, Quercus alba, Q. rubra, Carya spp., and flora, Persea borbonia, and Ilex opaca are commonly
Juglans nigra, all of these frequently on shallow, associated trees. Over large areas the small palm
rocky or sandy soils (‘glades’). It is also common in Sabal palmetto can dominate the understorey, but a
moister places near streams, especially in the west- more diverse shrub layer occurs especially on more
ern part of its range bordering the Great Plains, and stabilised sites. Past selective cutting of this variety
can grow well on a range from acid to neutral soils. (Southern Red-cedar) has led to dominance of oaks
Annual precipitation is less than for var. silicicola, and pines in many former ‘cedar’ forests. Further in-
with an effective summer drought limit of around land, var. silicicola becomes a minor component of
350 mm. pine forests and in some areas it is sympatric with
the more widespread var. virginiana. The climate is
470 subhumid to humid (1000–1600 mm of rain p.a.)
Conservation
with very mild winters.
IUCN: LC
Conservation
Juniperus virginiana L. var. silicicola (Small)
E. Murray, Kalmia 13: 8. 1983. Sabina silicicola IUCN: LC
Small, J. New York Bot. Gard. 24: 5. 1923; Juniperus
silicicola (Small) L. H. Bailey, Cult. Conif. N. Amer.:
Excluded names
18. 1933; Juniperus virginiana L. subsp. silicicola
(Small) E. Murray, Kalmia 13: 8. 1983. Type: USA: Juniperus gaussenii W. C. Cheng, Trav. Lab. Forest.
Florida, Dixie Co., Suwannee River, Hog Island Toulouse T. 1 (3, 8): 3. 1940.
[“shell mound south of mouth of Suwanee River”],
J. K. Small & G. K. Small 10030 (lectotype NY).
Taxonomic notes
W. C. Cheng (Wanjun Zheng) described Juniperus
Vernacular names
gaussenii when he was a research fellow at the Labo-
Southern red-cedar, Red-cedar, Sand cedar, Coastal ratoire Forestier de Toulouse, France working under
red-cedar, Coast juniper H. Gaussen. Its type, F. Ducloux 3928 (holo. P) is one
of several gatherings Ducloux made in Kunming
(Yunnan-sen) early in the 20th century. Cheng only
Description
cites one further collection, R. P. Maire s.n., coll. 10
Trees not conical with ascending branching habit Aug 1921, which had pollen cones and “fruit roux” so
when mature. Leaves small, 1.3–2 × 0.7–1 mm on perhaps indicating a monoecious plant (if gathered
ultimate branchlets, acute-acuminate, not apiculate. from a single tree). On this basis Cheng (op. cit.)
compared his new species with J. chinensis, to which
it is in his opinion very close. Differences cited are
Distribution
very slight though, and some, e.g. the colour of seed
SE USA: Alabama, Florida, Georgia, Louisiana, Mis- cones, an artifact resulting from comparison of very
sissippi, North Carolina, South Carolina (near the few specimens. Cheng had no experience with liv-
coast). ing plants of his species; had he seen them he might
TDWG codes: 78 ALA FLA GEO LOU MSI NCA SCA have concluded that there was much less affinity
with J. chinensis. Numerous plants in the Botanic
Garden of the Institute of Botany, Chinese Acad-
Ecology
emy of Sciences, Kunming (it is even used in hedges
Juniperus virginiana var. silicicola is restricted to a there), labeled as J. gaussenii, show characteristics
narrow coastal strip near sea level of the Atlantic in growth and habit, including foliage, suggesting
Ocean and the Mexican Gulf and the lowlands of affinity with J. squamata as well as with J. chinen-
central Florida. It is there associated with limestone sis. These plants strongly suggest a cultivar, which
and prehistoric shell middens, often bordering tidal fits well with the virtual absence of cones (retained
marshes or occurring in old dunes. Pinus elliottii, juvenile stage of leaves) and an uncommon crown
Quercus virginiana, Q. laurifolia, Magnolia grandi- shape not found in any true species. Verification of
herbarium ­collections in KUN and PE confirmed
Ecology
the cultivated status of J. gaussenii in all cases where
data on habitat were given. Serious doubts therefore Not known to occur in the wild and apparently pro-
arise concerning the status of this species, which is ducing very few seed cones, most of which may be
probably best regarded as a cultivar of possible hy- sterile.
brid origin.
Uses
Distribution
A highly ornamental shrub which can attain large
China: Yunnan, only known from cultivation in size; it is not known in cultivation outside China.
some numbers in and around Kunming and in Xi- 471
chou in SE Yunnan. Also present in other ancient
cultural centres in Yunnan, e.g. Dali.
 Keteleeria Carrière, Rev. Hort. 37: 449. 1866. Type: Keteleeria fortunei (A. Murray
bis) Carrière (Pinaceae).

Named after Jean-Baptiste Keteleer, a Belgian hor-

Taxonomic notes
ticulturist.
A total of 14 species and 1 variety have been de-
scribed in this genus, the majority of which were
Description
based on small differences in variable characters ob-
Monoecious evergreen trees with a monopodial served in few specimens or collections. In the first
472 trunk. Resin canals in wood (few), leaves and seed revision of the genus Keteleeria since Flous (1936),
cones. Banches in pseudo-whols, spreading more Farjon (1989) has reduced the number of species to
or less horizontally and ascending (Massart’s and/ the three accepted by most authors prior to 1936:
or Rauh’s model); capacity to coppice present. Bark K. davidiana (Bertr.) Beissner, K. evelyniana Mas-
scaly and often longitudinally fissured on lower part ters and K. fortunei (Andr. Murray) Carrière. This
of trunk. Terminal buds ovoid conical to subglo- taxonomy is now generally accepted, although in
bose, axillary lateral buds subglobose, with numer- Flora of China 4: 42–44 (1999) two more species,
ous, more or less triangular, imbricate and persistent K. hainanensis and K. pubescens, were maintained
bud scales without resin. Leaves spirally inserted (with some caveats).
and pectinately arranged, weakly dimorphic; leaves
on saplings, young trees and coppicing shoots rela-
Key to the species of Keteleeria
tively broad, lanceolate, thin and flat, with acute
acuminate apices and usually larger than leaves on It should be observed, that only the seed scales of the
mature trees, which are linear, narrower, thicker, central part of mature cones show these diagnostic
with more obtuse apices; all leaves with two primary characters fully; likewise, only shoots and leaves on
stomatal bands on abaxial side and a variable but branches of mature trees, not of (relatively) young
much lesser number or no stomata on adaxial side. plants or regrowth of coppiced plants, should be
Pollen cones in umbellate clusters of approximately compared using this key. Trees in cultivation often
5–10 from a single axillary bud, cylindrical, 1–1.5 cm tend to retain juvenile characters longer than is ob-
long; microsporophylls with 2 pollen sacs contain- served in natural habitats.
ing bisaccate pollen. Seed cones on leaved pedun-
cles, erect, oblong to cylindrical. Bracts relatively 1a. Seed scales of mature cones with convex,
small, sometimes trilobate and always cuspidate. rounded or truncate upper margin. Leaves
Seed scales persistent (cone rachis finally breaking short (less than 4 cm long). Shoots usually gla-
up after a few to several seasons), flabellate with a brous K. fortunei
relatively broad, pedicellate base. Seeds large, held 1b. Seed scales of mature cones with a more or less
in a deep cup, fully covered on one side and for a obtuse-acute, concave or recurved apex. Leaves
fourth on other side; membrane continuing in a often longer than 4 cm, but variable. Shoots
persistent, oblique wing 1.5–2.5 times length of seed. glabrous or pubescent 2
Germination hypogeal (unique in Pinaceae); seed- 2a. Seed scales equally wide as long or slightly
ling with 2–4 cotyledons. longer, lateral margins straight. Leaves with an
obtuse or truncate apex. Shoots brown pubes-
3 species. cent K. davidiana
2b. Seed scales longer than wide, lateral margins
usually concave. Leaves with a more or less
Distribution
mucronate apex. Shoots weakly pubescent or
Central and SE China, Vietnam, N Laos, Taiwan. glabrous K. evelyniana
Keteleeria davidiana (Bertrand) Beissn., Handb. grooved with resin vesicles, 10–15 × 6–8 mm, dull
Nadelholzk.: 424, f. 117. 1891. brown, on one side covered by the seed wing; seed
wing semi-trullate, 25–30 × 10–12 mm, lustrous light
brown.
Etymology
This species has been named after Armand David
Taxonomic notes
(1826–1900), a French Lazarist missionary to China
and an avid naturalist. Keteleeria davidiana is a highly variable species and
this has led several authors to describe new species
based on such aberrant specimens as seemed distinct.
Vernacular names
Most if not all of these have later been reduced to 473
tijian yushan (Chinese) synonymy or, at most, to varieties of this widespread
species. In Farjon (1989, 1990) K. formosana was re-
duced to synonymy, but in Flora of China 4: 44 (1999)
Description
it has been retained as a variety, based on a single and
Trees to 40–50 m tall, d.b.h. to 2–2.5 m; trunk mo- rather obscure distinction. As a clearly disjunct ele-
nopodial; bark rough and scaly, fissured in lower ment within the species, endemic to Taiwan, it may
part of trunk, dark grey brown. Branches of first and be appropriate to give it the benefit of the doubt as a
second order long, heavy, spreading and ascending; mere variety of K. davidiana, as is done here.
crown broad conical or domed, often open in old
trees. Branchlets slender, firm, (light) reddish brown
Distribution
or yellowish brown, becoming grey; surface ridged
and grooved; young shoots usually densely brown China: NE Yunnan, SE Sichuan, Chongqing, SE
pubescent, but soon glabrous, leaf scars small, cir- Gansu, S Shaanxi, NW Guizhou, SW Hubei, SW
cular. Vegetative buds ovoid globose, 3–5 × 2–4 Hunan, N Guangxi; Taiwan.
mm, not resinous; bud scales triangular, obtuse, TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
appressed, persisting several years. Leaves directed SC CHC-YN CHN-GS CHN-SA CHS-GX CHS-HN 38
forward, (1.5–)2–5(–5.5) cm long, 2.5–4.5 mm wide, TAI
slightly twisted and/or narrowed at base, narrowly
linear, ligulate linear, or lanceolate in young plants,
Ecology
flattened, with slightly recurved margins, obtuse or
truncate (in young plants acute) at apex, with a lon- Keteleeria davidiana occurs from hills to low moun-
gitudinal midrib on both surfaces; stomata none or a tains throughout much of E China, at elevations of
few near apex above, in two broad bands below; leaf (300–)600–1000(–1300) m a.s.l. It grows on the red
colour (glaucous) green above, greenish white sto- and yellow earth, which are acid, podzolic soils poor
matal bands below. Pollen cones pedunculate, 1–1.5 in nutrients, or on brown forest soils. The climate is
cm long, yellow with brown perular scales. Seed humid, continental warm temperate to subtropical,
cones lateral or (sub)terminal, solitary or paired; pe- with annual precipitation between 1000 and 2000
duncles 1.5–6 cm long, leaved as shoots, at an angle mm. It is a constituent of the mixed mesophytic
to axis of cone; cones short or long cylindrical, with forest formation (Wang, 1961), together with many
obtuse apex, (5–)8–21 cm long, 3.5–6 cm wide with genera and species of broad-leaved deciduous trees,
opened scales, ripening to light or dark brown; cone and some other gymnosperms, such as Pinus mas-
rachis deciduous with cone, or slowly disintegrat- soniana, P. bungeana, Cunninghamia lanceolata,
ing, narrowly conical. Seed scales subcordate, with Cupressus funebris, Torreya grandis, and Podocar-
often reflexed apical end or margin, 2.6–3.2 × 2.2–2.8 pus nakaii (Taiwan). It also occupies the evergreen
cm at mid-cone; abaxial surface striated, sometimes broad-leaved forest formation (Guizhou, Taiwan),
pubescent in young cones, but soon glabrous; upper with numerous (sclerophyllous) evergreen tree spe-
margin erose denticulate in young cones, becoming cies and Pinus spp. It rarely forms pure stands. It oc-
entire. Bracts narrowly spathulate, with cuspidate curs in the parts of China where deforestation has
or tridentate apex, 1.5–2 cm long, straight, slightly been going on for millennia, leaving very little of the
exserted with opened seed scales. Seeds oblong, primeval forest. Keteleeria survives coppicing and,
 like many species of Pinus, appears to act as a pio- TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
neer in secondary vegetation. SC CHC-YN CHN-GS CHN-SA CHS-GX CHS-HN

Uses Conservation
The timber of this species is used for construction, IUCN: LC
carpentry and firewood. It is commonly planted in
China as an amenity tree in parks, near temples and Keteleeria davidiana (Bertrand) Beissn. var. for-
sometimes as a street tree. It was introduced to Eu- mosana (Hayata) Hayata, J. Coll. Sci. Imp. Univ.
rope on several occasions and is the most common- Tokyo 25 (19): 221, f. 11. 1908. Keteleeria formosana
474 ly grown species in arboreta and botanic gardens. Hayata, Gard. Chron., ser. 3, 43: 194. 1908; Kete­
Outside tree collections in Europe and the USA it leeria davidiana (Bertrand) Beissn. subsp. formo­
is very rarely found growing, even though, under sana (Hayata) E. Murray, Kalmia 12: 21. 1982. Type:
good conditions, it can be grown easily from cut- Taiwan: locality unknown, N. Konishi s.n. (holotype
tings. This species, which seems to be more hardy BM).
than the others, should receive more attention from
dendrologists and arboriculturists in parts of the
Description
world with mild winters, now apparently expanding
northwards at least in Europe and North America. Leaves 2–4 cm long, 3–4 mm wide; leaf scars pro-
truding on branches, dark.
2 varieties are recognized:
Distribution
Keteleeria davidiana (Bertrand) Beissn. var.
davidiana. Pseudotsuga davidiana Bertrand, Bull. Taiwan (Taipei Co., Taitung Co.).
Soc. Philom. Paris, sér. 6, 9: 38. 1872. Type: China: TDWG codes: 38 TAI
Sichuan, Lunganfu [“Longan-fou”], A. David 36
(holotype P). Fig. 151, 152
Conservation
Keteleeria calcarea W. C. Cheng & L. K. Fu, Acta IUCN: EN (C2a)
Phytotax. Sin. 13 (4): 82. 1975; Keteleeria davidiana
(Bertrand) Beissn. var. calcarea (W. C. Cheng &
L. K. Fu) Silba, Phytologia 68: 34. 1990. Keteleeria evelyniana Mast., Gard. Chron., ser. 3,
Keteleeria pubescens W. C. Cheng & L. K. Fu, Acta 33: 194. 1903. Type: China: Yunnan, Jiangchuan,
Phytotax. Sin. 13 (4): 82. 1975; Keteleeria davidiana A. Henry 11815 (holotype NY).
(Bertrand) Beissn. var. pubescens (W. C. Cheng &
L. K. Fu) Silba, Phytologia 68: 34. 1990. Keteleeria hainanensis Chun & Tsiang, Acta Phyto-
Keteleeria xerophila J. R. Xue & S. H. Hao, Acta Bot. tax. Sin. 8 (3): 259. 1963; Keteleeria evelyniana Mast.
Yunnanica 3 (2): 249–250, f. 1–5. 1981; Keteleeria var. hainanensis (Chun & Tsiang) Silba, Phytologia
fortunei (A. Murray bis) Carrière var. xerophila 68: 35. 1990.
(J. R. Xue & S. H. Hao) Silba, Phytologia 68: 36. 1990. Keteleeria evelyniana Mast. var. pendula J. R. Xue,
Acta Phytotax. Sin. 21 (3): 253. 1983.
Description
Etymology
Leaves (1.5–)2–5(–5.5) cm long, 2.5–4.5 mm wide;
leaf scars obscurely protruding on branches. This species commemorates the English arboricul-
turist John Evelyn.
Distribution
Vernacular names
China: Chongqing, SE Gansu, N Guangxi, Guizhou,
SW Hubei, SW Hunan, S Shaanxi, Sichuan, NE Yun- Yunnan yushan (Chinese); Du sam nui dat, Ngo
nan. tung (Vietnamese)
 (the others are species of Pinus). It is found in
Description
mountainous areas in SE Asia at elevations of 700 to
Trees to 30–40 m tall, d.b.h. to 1–1.5 m; trunk mo- 2700 m a.s.l. (to 3000 m according to Wilson, 1926),
nopodial, usually straight; bark becoming rough and but generally not above 2000 m. The soil is mainly
scaly, dark grey brown. Branches of first and second red earth (in China and Laos); the climate is humid,
order long, curved, spreading, ascending near the top; tropical to temperate at high altitudes, often with
crown (broad) conical, irregular in old trees. Branch- more than 2000 mm precipitation annually. It is a
lets slender, firm, (light) reddish brown or yellowish minor constituent of the evergreen broad-leaved for-
brown, turning grey brown; pubescence on young est formation, which occurs in mountains above the
shoots only, weak or absent; leaf scars small, circular. tropical lowland rainforest. In Yunnan and N Laos,
Vegetative buds ovoid globose or ovoid conical, 4–6 it also occurs in mixed evergreen oak forest, with 475
× 3–4 mm, not resinous; bud scales triangular, ob- Cunninghamia lanceolata (Yunnan), Podo­carpus
tuse, appressed, dull brown, persisting several years. spp., Cephalotaxus fortunei, Fagaceae, Lauraceae,
Leaves usually pectinate, on terminal shoots some- Magnoliaceae, etc.
times assurgent, directed forward, (2–)3–6.5(–8) cm
long, 2–4 mm wide, slightly twisted or only narrowed
Conservation
at base, linear, often falcate, lanceolate in young or
coppiced plants, flattened; apex usually mucronate, In Vietnam, this species has been assessed as Vul-
sometimes obtuse (acute in young plants); stomata nerable (VU, criteria A2cd), because it has been
usually in several lines near central rib above, in two over-exploited for local use of the timber and large
broad bands separated by a midrib below; leaf colour parts of its natural habitat have been converted to
(glaucous) light or dark green above, greenish white agriculture. Some stands of this tree are protected
stomatal bands below. Pollen cones pedunculate, within protected areas and, officially at least, exploi-
1–1.5 cm long, yellow, with brown perular scales. Seed tation is now limited by government forest policy.
cones lateral, usually solitary, erect; peduncles at an The overall situation in China cannot be much dif-
angle to cone axis, 2–6 cm long, leaved as shoots; cone ferent, but at least in parts of Sichuan and Yunnan
long cylindrical (when fully developed!), with obtuse K. evelyniana is widespread and still more common.
apex, (4–)9–20(–25) cm long, (3–)4–6.5(–9?) cm The conservation status in Laos is unknown.
wide with opened scales, ripening to (light) brown, IUCN: LC
often lustrous; cone rachis narrowly conical. Seed
scales subcordate oblong, with narrowed apex and
Uses
more or less concave margins, straight or recurved,
3–4 × 2.5–3 cm at mid-cone; abaxial surface striated, The timber of this species is used locally for con-
glabrous; upper margin erose denticulate in young struction purposes and firewood. It has been intro-
cones, later entire or erose. Bracts ligulate spathu- duced to the UK by George Forrest and to the USA
late, with cuspidate or trilobate apex, 1–1.5 cm long, (California) by Joseph Rock, but until recently it re-
straight, slightly exserted with opened seed scales. mained a rarity in arboreta and botanic gardens. It is
Seeds oblong, grooved with resin vesicles, 9–14 × 5–7 now quite commonly planted in the USA. Its exten-
mm, dull brown, on one side covered by the seed sive N-S range indicates growing conditions from
wing; seed wing semi trullate, 20–30 × 12–15 mm, mild temperate to near tropical; with the universal
lustrous yellowish brown. requirement of warm, moist summers.

Distribution
Keteleeria fortunei (A. Murray bis) Carrière, Rev.
China: Hainan Island, SW Sichuan, Yunnan; Laos;
Hort. 37: 449. 1866. Picea fortunei A. Murray bis,
Vietnam.
Proc. Roy. Hort. Soc. London 1862 (2): 419–425,
TDWG codes: 36 CHC-SC CHC-YN CHH 41 LAO VIE
f. 85–97. 1862. Type: China: Fujian, Min River,
Fuzhou, R. Fortune 52 (lectotype BM). Pl. 19
Ecology
Keteleeria evelyniana is one of the few species of Keteleeria oblonga W. C. Cheng & L. K. Fu, Acta
Pinaceae occurring in near tropical environments Phytotax. Sin. 13 (4): 82. 1975; Keteleeria fortunei
 (A. Murray bis) Carrière var. oblonga (W. C. Cheng vesicles, 10–13 × 5–6 mm, dull brown, on one side
& L. K. Fu) L. K. Fu & Nan Li, Novon 7 (3): 261. 1997. covered by the seed wing; seed wings more or less
cuneate, with oblique end, 13–20 × 8–12 mm, yellow-
ish brown.
Etymology
This species was named after Robert Fortune (1812–
Distribution
1880), an early English collector of plants in China.
China: Fujian, N Guangdong, Guangxi, Guizhou,
Hong Kong, S Hunan, SW Jiangxi, SE Yunnan, Zhe-
Vernacular names
jiang.
476 yushan (Chinese) TDWG codes: 36 CHC-GZ CHC-YN CHS-FJ CHS-GD
CHS-GX CHS-HK CHS-HN CHS-JX CHS-ZJ
Description
Ecology
Trees to 30 m tall, d.b.h. to 1–1.5 m; trunk monopo-
dial, straight, often short and branching low; bark Keteleeria fortunei occurs in the hills or low moun-
becoming thick and fissured, dark grey brown. tains of SE China, in the “red and yellow earth re-
Branches of first order heavy, long, spreading wide; gion” (Wang, 1961), at elevations between 380 and
crown broad, often dome shaped. Branchlets slen- 1200 m a.s.l. The climate is humid, warm temper-
der, firm, (light) reddish brown or yellowish brown, ate to subtropical, with annual precipitation be-
glabrous, or rarely with some short hairs in grooves; tween 1300 and 2000 mm. It occurs in two forest
leaf scars small, circular. Vegetative buds ovoid coni- formations: the mixed mesophytic forest, and, more
cal or subglobose, 3–5 × 2–4 mm, not resinous; bud usually, the evergreen broad-leaved forest. Besides
scales triangular, obtuse and appressed, persist- many angiosperm trees, such as evergreen sclero-
ing several years. Leaves spreading at 45–90° from phyllous oaks and lauraceous trees, a few additional
shoot, (1.2–)1.5–3(–4) cm long, 2–4 mm wide, slight- gymnosperms are also found in the latter formation:
ly twisted and narrowed at base, narrowly linear Pseudotsuga sinensis, Cryptomeria japonica, Cepha-
to ligulate, flattened, with a raised midrib on both lotaxus fortunei, and Taxus chinensis.
surfaces; apex obtuse, rarely somewhat acute (but
acute on young or coppiced plants); stomata absent
Conservation
or a few near apex above, in two broad bands sepa-
rated by the midrib below; leaf colour green, green- Although relatively widespread in distribution, cen-
ish white below. Pollen cones peduncled, 1–1.5 cm turies of deforestation in southern China have un-
long, yellow, with brown perular scales. Seed cones doubtedly reduced the forests in which this species
lateral or (sub)terminal, erect; peduncles 2–5 cm naturally occurs. It is, however, capable of regenera-
long, leaved; shape cylindrical, with obtuse apex, tion in secondary vegetation; a decline in primary
6–18 cm long, 3.5–6.5 cm wide with opened scales, forest can therefore not be directly translated into a
ripening to (greyish) brown. Seed scales subcordate decline of the species.
orbicular, with convex, rounded or nearly straight IUCN: NT
upper margin, 1.8 3.2 × 2 3.3 cm at mid-cone; surface
smooth, usually striated longitudinally, in imma-
Uses
ture cones often puberulent, soon glabrous; upper
margin erose denticulate in young cones, later finely The wood of this species is used locally for construc-
denticulate or entire. Bracts ligulate spathulate; apex tion and firewood. It is quite commonly planted in
cuspidate, sometimes weakly trilobate, 1–1.5 cm China, but rare in cultivation elsewhere; it was in-
long, included or slightly exserted, visible when seed troduced to England from seed collected in Hong
scales are opened. Seeds oblong, grooved with resin Kong.
 9
477

4
2

Pl ate 19 . Keteleeria fortunei. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone. 4. Immature seed
cone. 5. Seed scale with seeds. 6. Seeds. 7. Leaves. 8. Leaf apices. 9. Pollen cones.
 Lagarostrobos Quinn, Austral. J. Bot. 30 (3): 316. 1982. Type: Lagarostrobos franklinii
(Hook. f.) Quinn [Dacrydium franklinii Hook. f.] (Podocarpaceae).

Greek: lagaros = narrow; strobilos = cone. in appearance, 1–1.5 × 1 mm, keeled abaxially; apex
obtuse. Leaves amphistomatic, stomata conspicu-
ous, scattered. Pollen cones terminal, sessile, 4–6
Description
mm long, 2–2.5 mm wide; microsporophylls 10–15
See the species description. (–20), rhombic to triangular, with minutely denticu-
late upper margins and with two basal pollen sacs
478 containing trisaccate pollen. Seed cones terminal on
Distribution
decurved short branchlets, 4–5 mm long, consisting
As for the species. of 5–8(–10) spirally arranged fertile bracts, each with
a single erect ovule on the adaxial side. Seeds up to
5–8 per cone, usually fewer, morphologically erect
Lagarostrobos franklinii (Hook. f.) Quinn, Austral. but topographically pendent, ca. 2.2 × 2 mm, dor-
J. Bot. 30 (3): 316. 1982. Dacrydium franklinii Hook. siventrally compressed to nearly rounded in cross-
f., London J. Bot. 4: 152, t. 6. 1845. Type: Australia: section, notched ar apex, light brown, enclosed at
Tasmania, Huon River, [“grows at Mcq Harb”], base in a dry, papery epimatium.
A. Cunningham s.n. (a-c) (lectotype K). Fig. 153, 154
Distribution
Etymology
Australia: Tasmania (mainly S & W parts, along riv-
The species epithet commemorates Sir John Frank- ers).
lin, a Governor of Tasmania in the early years of the TDWG codes: 50 TAS
colony.
Ecology
Vernacular names
Lagarostrobos franklinii is mostly a riparian spe-
Huon pine cies, usually on river banks or close to rivers, but
occasionally occurring on wet hill sides away from
main water courses in temperate rainforest. It forms
Description
groves dominated by Huon pine, marking stream
Evergreen, predominantly dioecious trees to 25 m courses at low altitudes from sea level to 150 m, and
or perhaps 30 m tall; trunk to 1.5(–2) m d.b.h. (few it grows on some hills to 750 m in the mountainous
trees of this size now exist). Bark becoming longi- west country. In a few areas it is observed to have
tudinally fissured, fibrous, exfoliating in scales and spread by layering, most notably on Mt. Read, where
strips, ultimately 4–6 cm thick, grey-brown. Crown several hectares are believed to be occupied by stems
of young trees more or less conical or pyramidal, forming a single clone. It is often accompanied by
of old mature trees spreading, with large ascending Nothofagus cunninghamii, Eucryphia lucida, Ano­
main branches. Frequently layering, sometimes ex- pterus glandulosus and ferns, with Eucalyptus obli-
tending over large areas. Foliage branchlets slender, qua growing nearby on higher ground.
1–1.2 mm diam. including scale leaves, long on seed-
lings and young plants, or on shaded, pendulous
Conservation
branches, short and more rigidly spreading on sun-
exposed branches in the crown of mature trees. Ju- This species is not threatened under current (2001)
venile leaves on seedlings and young plants, spirally IUCN criteria; it was listed as LRcd in the Conifer
arranged, decurrent, spreading to all sides with free Action Plan (Farjon & Page, 1999), but the subcate-
apex, 1–2 mm long, keeled abaxially, concave adaxi- gory “conservation dependent” is no longer recog-
ally; apex curved forward, acute. Adult leaves spi- nized. An estimated 15% of its habitat has been lost
rally arranged, imbricate and appressed, rhomboid through inundation for hydroelectric schemes and
to fire over the past 100 years or so. Extensive log- and resources of good timber trees were exhausted.
ging in the past has removed nearly all large trees, The timber was exported in the colonial period of
but there is regrowth nearly everywhere; ca. 85% of the 19th century and convicts were employed in the
the remaining area of occupancy (AOO) of Huon logging and transport operations by river. Its wood
pine is now protected in reserves. One stand of the is hard and durable and was mainly used for boat
species has been made available for access to craft building and decks of sailing ships. Today, virtually
wood from dead and downed timber, but there is all large trees have gone, but a few escaped the log-
no cutting of living Huon pine allowed. Fire man- gers and are now assidiously protected. A limited
agement appears to be the main priority at present amount of down and dead wood is now used for
to ensure its continued conservation under the cur- wood crafts and cabinet work. The species is rare in
rently prevailing policies. cultivation, but present in a few botanical collections 479
IUCN: NT and arboreta. This tree grows very slow, both in the
wild and in cultivation, and is tolerant of light frost.
Uses
Huon pine was once the most important timber tree
of Tasmania, but its exploitation was unsustainable
 Larix Mill., Gard. Dict., Abridg. Ed. 4, vol. 1. 1754. Type: Larix decidua Mill.
(Pinaceae).

Larix is the classical Latin name for larches. and, to a lesser degree, in the leaves. These groups
have been formalized in sections:
–– Section Larix with species L. decidua (type),
Description
L. sibirica, L. gmelinii, L. ×czekanowskii, L. lar-
Monoecious, deciduous trees with a monopodial icina and L. kaempferi
trunk. Resin canals in wood, leaves and seed cones. –– Section Multiserialis Patschke with species
480 Branches at regular intervals on trunk, spreading L. griffithii (lectotype), L. potaninii, L. lyallii,
and assurging or highest order branches drooping L. occidentalis and L. mastersiana
to pendulous (Rauh’s model). Bark thick, scaly and
fissured, with large plates in some species. Shoot This classification was adopted im my book Pina­
dimorphism pronounced; lateral short shoots give ceae (Farjon, 1990); but since then a number of pa-
rise to leaves, long shoots and reproductive or- pers have been published attempting to reconstruct
gans. Leaves narrowly linear, (sub-)flexible, obtuse a phylogeny of the genus. One of these, based on
to acutish at apex, more or less flattened or broad morphological data and including evidence from
triangular, sometimes diamond shaped in cross sec- the fossil record (LePage & Basinger, 1995) divides
tion, hypo- or amphistomatic. Pollen cones solitary the genus in two groups: one with short bracts and
at apex of short shoots and often numerous; micro- one with long bracts. This is a similar classification
sporophylls with 2 pollen sacs containing globular, as the one above, but with L. kaempferi transferred
smooth pollen with a narrow equatorial ridge. Seed to section Multiserialis; the fossils are all placed with
cones solitary at apex of short shoots on a curved the other, short-bracted group. The results of two
peduncle, more or less erect, persistent, falling at- analyses of molecular (DNA sequence) data appear
tached to branches. Bracts short and hidden or long to give results that group species according to their
and exserted. Seed scales rounded or emarginate, geographical distribution when based on plastid
more or less convex, with a short pedicellate base DNA (Wei & Wang, 2003) and nuclear ribosomal
and persistent. Seeds ovoid, held in a shallow cup (ITS) DNA (Wei & Wang, 2004). Here, there is a
covering one side of the seed, which extends in a North American group (L. occidentalis, L. laricina), a
relatively short, persistent wing. Seedlings with 5–7 North Eurasian group and a Sino-Himalayan group,
(usually 6) cotyledons. but with L. sibirica ‘behaving’ aberrantly in the cp-
DNA based analysis and L. lyallii (North America)
11 species. not represented in the analyses. In another study of
this kind (Semerikov et al., 2003) conflicting phyl-
ogenies were inferred from cpDNA and ITS (with
Distribution
L. lyallii included). The only consistently congruent
North America: central Alaska (disjunct); from the result seems to be a clade with the North Ameri-
NW Territories to Newfoundland; northern Rocky can species L. laricina, L. lyallii and L. occidentalis.
Mountains and Cascade Range; New England states These belong to two different traditional sections.
(USA). Eurasia: Europe (Alps and Carpathians); It appears therefore, that at present a classification
NE Russia across Siberia to Kamchatka and Sakha- informed by a robust phylogeny that would help to
lin; Japan (disjunct), NE China; Sino-Himalayan explain character transformations in an evolution-
mountain system. ary context still eludes us. No formal classification is
therefore presented here.
Synopsis
Key to the species of Larix
The genus Larix has been divided into two groups
based on morphological characters in the seed cones 1a. Bracts of mature seed cones shorter than seed
scales, usually barely visible in opened cones;
 ripe seed cones globose, ovoid or ovoid-coni- Larix ×czekanowskii Szafer, Kosmos 38: 1297. 1913.
cal. Leaves carinate on invers-dorsal side (be- Type: Russia: Siberia, Tunguska River, below mouth
low) only 2 of Tomezoy River, A. Czekanovsky & F. Müller s.n.
1b. Bracts of mature seed cones (much) longer than (lectotype LE).
seed scales, straight or reflexed; ripe seed cones
ovoid-conical to oblong-cylindrical. Leaves
Etymology
carinate on both sides, rarely only on invers-
dorsal side 7 This nothospecies was named after Aleksander
2a. Seed scales concavo-convex, with strongly re- P. Czekanowski (1833–1876), a Polish geologist who
curved upper margin; seed cones broad ovoid, studied the geology of the Lake Baikal Basin while
(1.5–)2–3(–3.5) cm long L. kaempferi in exile. 481
2b. Seed scales convex or more or less straight, with
flat upper margin; seed cones variously shaped
Vernacular names
3
3a. Seed scales 10–20 per cone, not spreading; seed No common names have been recorded for this
cones 1–2 cm long L. laricina taxon.
3b. Seed scales (15–)20–40(–45) per cone, spread-
ing; seed cones (1.5–)1.8–5(–6) cm long 4
Description
4a. Leaves hypostomatic (all or nearly all stomata
on one side) L. gmelinii Trees which are ‘intermediate’ between Larix sibir-
4b. Leaves amphistomatic (stomata on both sides, ica and L. gmelinii and appear to be common in
but usually more on one side) 5 a wide belt stretching N to NW from Lake Baikal
5a. Seed scales of green cones (densely) pubescent, share some character states of both species, par-
becoming more glabrous with age L. sibirica ticularly evident in the cones. The seed cones are
5b. Seed scales of green cones always glabrous 6 smaller than those of L. sibirica and resemble those
6a. Seed scales incurved, straight or slightly re- of a large-cone variety of L. gmelinii: var. principis-­
curved; upper margin repand or emarginate, rupprechtii; the seed scales are less pubescent than
sometimes entire (European species) those of L. sibirica. They have been interpreted by
L. decidua Russian botanists and foresters as of natural hybrid
6b. Seed scales incurved; upper margin entire (Si- origin.
berian species) L. ×czekanowskii
7a. Apex of bracts in seed cones broadly acute
Distribution
L. potaninii
7b. Apex of bracts in seed cones cuspidate, often Central Siberia: from Lake Baikal to the mouth of
narrow and elongated 8 the Yenisei River
8a. Mature seed cones 5–8(–11) cm long, cylindri- TDWG codes: 30 IRK KRA
cal L. griffithii
8b. Mature seed cones 2.5–5(–6) cm long, variously
Ecology
shaped but longer than wide 9
9a. Leaves amphistomatic. Bracts straight, with an Larix ×czekanowskii occurs in central Siberia, where
abruptly narrowing cusp L. lyallii it forms taiga forest with Picea obovata, Pinus sylves-
9b. Leaves hypostomatic. Bracts recurved (at least tris and broad-leaved trees such as Betula pendula
the cusp) 10 and Populus spp., broadly following the Yenissei
10a. Seed scales recurved, mostly emarginate River. It grows on a great variety of soils, from peat
(sometimes entire); bracts with a thin, elon- bogs to well drained, sandy or rocky soils, where
gated cusp L. occidentalis it has its optimum. The climate is very cold (min.
10b. Seed scales more or less straight, entire or temp. –55 °C), continental or subarctic, dry, with
slightly emarginate; bracts short cuspidate very long winters.
L. mastersiana
 by a keel; leaf colour light green, darkening, yellow
Conservation
in autumn. Pollen cones terminal on short shoots,
IUCN: LC numerous on pendulous long twigs, 0.5–1 cm long,
yellow, perular scales with fimbriate margins, red-
dish. Seed cones terminal on short shoots, turning
Uses
erect; peduncles curved, 0.5–1 cm long, subtended
The wood of this larch is durable and used in con- by leaves; cones ovoid or ovoid oblong, with obtuse
struction, traditionally for log houses in Siberia for apex, (1.5–)2.5–4(–6) cm long, (1.2–)1.5–3(–3.5) cm
which the wood is roughly hewn to shape, but un- wide with opened scales; colour (immature) dark
treated. It has been widely used for railroad sleepers red or purplish, sometimes green, maturing to pale
482 e.g. on the famous Trans Siberian Railroad. Larch green with purplish margins of seed scales, ripen-
wood is also milled for construction timber and ve- ing to (dark) brown, old cones grey. Seed scales
neer, and pulped for the paper industry. This natural 25–35, ovate to suborbicular, slightly convex, 7–15 ×
hybrid larch is not known to be in cultivation, but it 6–13 mm; surface striated, shining smooth in older
may be present in e.g. Scandinavia under the name cones, reddish pubescent near base, later glabrous;
L. sibirica. upper margin entire, incurved, repand or emargin-
ate, sometimes slightly recurved; base narrowed.
Bracts ligulate, with acicular or cuspidate apex,
Larix decidua Mill., Gard. Dict., ed. 8: Larix No. 1. length ⅓-½ × seed scales, mostly included, but vis-
1768. ible with opened scales. Seeds ovoid-cuneate, 4 × 2.5
mm, dark brown-grey; seed wings oval, 6–10 × 4–6
mm, light brown.
Etymology
The species epithet refers to the deciduous (season-
Distribution
ally falling) leaves.
Europe: Alps, Carpathians, Slovenian Mts., S Po-
land.
Vernacular names
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SK
European larch; Gemeine Lärche (German); Mélèze GER POL SWI 12 FRA-FR 13 ITA-IT ROM YUG-SL 14
d’Europe (French) UKR-MO UKR-UK

Description Ecology
Trees to 40–50(–55) m tall, d.b.h. to 1.5–2.5 m; trunk Larix decidua occurs in the high mountains of
monopodial, straight or curved at base; bark deeply central Europe, at altitudes between (600–)1000–
fissured, breaking into large plates, flaking, exposing 2200(2500) m a.s.l., in the Central Alps it usually
reddish inner bark. Branches heavy, long, curved forms the tree limit. The soils are neutral to acidic,
down, ascending near the ends; branches of second mostly on granitic rock. The climate has cool, moist
order long, slender, pendulous; crown pyramidal in summers and cold, snowy winters, but annual pre-
young trees, widening in old trees. Branchlets thin, cipitation rarely exceeds 1000 mm. Pure stands are
slender, flexible, or stout, pink buff or (pale) yel- uncommon, more often it is mixed with Pinus cem-
lowish; glabrous, or slightly pubescent when young; bra in the Alps, below 1800 m also with Picea abies.
short shoots cylindrical or subglobose, 0.3–1 cm
long. Vegetative buds ovoid, 3 × 2 mm, not resin-
Uses
ous. Leaves on long shoots remote, appressed for-
ward against the shoot or spreading; on short shoots The wood of European larch is valued for its durabil-
crowded in false whorls, 30–40, (2.3–3.5(–4) cm long, ity and has been used for centuries in the Alps and
0.5–1 mm wide, narrowly linear, soft, flexible, in Carpathians to build houses; traditional style houses
cross-section broad triangular or flattened, carinate still use well seasoned wood of this species. Other
below, obtuse or acutish at apex; amphistomatic, sto- traditional uses are fences, gates, feeding racks, and
mata on lower side in two narrow bands separated water troughs for animals. Due to its durability the
wood of European larch has been used extensively
Conservation
for railway sleepers, until these were replaced by
concrete and iron structures in modern times. Trees IUCN: LC
with a curved base were split and hollowed and the
two halves joined to make ‘Alphorns’, large wind Larix decidua Mill. var. polonica (Racib. ex
instruments with a far carrying low tone; competi- Wóycicki) Ostenf. & Syrach, Pflanzenareale 2: 63.
tions to blow the horn are still held in some regions 1930. Larix polonica Racib. ex Wóycicki, Obraz.
of the Alps. This species has been introduced in the Rosl. Król. Polsk. 2: 15–16, t. 1. 1912. Type not
lowlands of Europe for plantation forestry as well as designated.
an amenity tree. In horticulture for gardens it is not
so common, although a modest number of cultivars 483
Description
is known, most with various branching habits.
Long shoots pale yellowish; leaves flattened. Seed
3 varieties are recognized: cones ca. 1.5 × 1.2 cm, with suborbicular, convex
scales with a round, entire upper margin.
Larix decidua Mill. var. decidua. Type: Illustration
“Larix folio deciduo Conifera” in Miller, Cat. Pl.: 43,
Distribution
t. 11. 1730 (lectotype). Fig. 155, 156, 157
Poland (headwaters of Wista River, Carpathians?).
TDWG codes: 11 POL
Description
Seed cones 2.5–4.5 cm long, 1.5–3 cm wide when
Ecology
opened; seed scales with a rounded or more or less
repand to emarginate upper margin. This variety has been found in scattered stands from
the slopes of the Tatra Mountains at ca. 1200 m a.s.l.
down to the plains at ca. 150 m a.s.l. It is commonly
Distribution
associated with Pinus sylvestris and Betula pendula,
Europe: Alps, Carpathians, Slovenian Mts. sometimes with Quercus robur.
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SK
GER SWI 12 FRA-FR 13 ITA-IT YUG-SL
Conservation
In forests surrounding the headwaters of the Wista
Conservation
River in Poland, larches have become very rare. In
IUCN: LC most areas only ancient relict trees survive and there
is little or no successful regeneration. This could
Larix decidua Mill. var. carpatica Domin, Sborn. partly be the result of historical climate change,
Vyzk. Ustav Zemed. R. Č. S. 65: 149. 1930. Type: helped by forest management practice and land uses
Slovakia, Tatra Mts., V. Krajina s.n., 7 sep 1931 that have favoured pioneer broad-leaved trees and
(holotype PR). pines. The geographical extent of this variety is un-
certain and could be greater than presently known.
IUCN: VU (B1+2c)
Description
Seed cones 4–6 cm long, 2.5–3.5 cm wide when
opened; seed scales often with an entire, rounded Larix gmelinii (Rupr.) Kuzen., Trudy Bot. Muz.
upper margin. Rissijsk. Akad. Nauk 18: 41. 1920.

Distribution Etymology
East Europe: Carpathians. This species was named after Johann Georg Gmelin
TDWG codes: 13 ROM 14 UKR-MO UKR-UK (1709–1755), a botanist who traveled widely in Sibe-
ria.
 Vernacular names Distribution
Dahurian larch; Listvennitsa daurskaya (Russian); NE China: Hebei, Manchuria, Nei Monggol [Inner
luoye song (Chinese); Gui-natsu (Japanese) Mongolia], N Shanxi; North Korea; NE Mongolia;
Russian Federation: E Siberia, Russian Far East.
TDWG codes: 30 BRY CTA IRK YAK 31 AMU KAM
Description
KHA KUR MAG PRM SAK 36 CHI-NM CHM CHN-HB
Trees to 30–35 m tall, d.b.h. to 1–1.5 m; trunk mo- CHN-SX 37 MON 38 KOR-NK
nopodial; bark on trunk dark red brown, with
greyish plates, finely scaly. Branches long, spread-
Ecology
484 ing horizontally, ascending near the top; branches
of second order relatively short, firm, drooping, Larix gmelinii occupies a very large area and there-
not pendulous; crown broad pyramidal, open, flat fore occurs in a wide range of habitats: on lowland
topped and irregular in old trees; Branchlets thin, subarctic plains, in river valleys, in mountains and
slender, light pink brown to orange-brown or dark also on the edges of moors and swamps. Its alti-
purplish brown; pubescence variable, weak or tudinal range is between 300 m and 1800 m a.s.l.
dense, or glabrous; short shoots small, 3–6 mm, cy- The soils are affected by permafrost over much of
lindrical. Vegetative terminal buds ovoid globose, the range; the climate in large parts of its range is
with acute apex, not or slightly resinous; bud scales continental subarctic, with very cold winters, and
ovate or obtuse triangular, dark brown or purplish relatively dry (400 mm to 500 mm in the Greater
brown. Leaves on short shoots crowded in false Hinggan Range) to extremely dry in the subarctic
whorls, 20–30(–35), 1.5–3(–4) cm long, 0.5–0.8 mm interior. In more maritime areas of the Russian Far
wide, narrowly linear, widest near apex, soft and East precipitation rises to 1000 mm and more. It is
flexible, cross-section diamond shaped, or flattened the only tree species in E Siberia reaching the tree
and keeled below; apex obtuse or acutish; stomata in limit at 70° N, but in the more southern part of its
two narrow bands separated by a keel below, none range it is commonly mixed with Abies sibirica, Pi-
or a few above; leaf colour bright green, yellow in cea obovata and Pinus sylvestris. In the boreal conif-
autumn. Pollen cones terminal on short shoots, erous forest it is a climax species only on permafrost
5–7 mm long, yellow. Seed cones terminal on short or peaty soils. In the high swamps and bogs in Sa-
shoots, turned erect; peduncles short, curved; cones khalin and on the Kuril Islands it usually forms pure
ovoid or ovoid-oblong, widening with opened stands, but on somewhat drier sites it is mixed with
scales; apex truncate, 0.8–3.5(–4.5) cm long, 0.8–2 Abies sachalinensis var. sachalinensis, Picea jezoensis,
(–3) cm wide with opened scales; colour (imma- Alnus hirsuta, Betula japonica, B. ermanii, and Salix
ture) purplish red or sometimes light green, matur- spp. In central Kamchatka occurs a relict taiga forest
ing to rose-purple or greenish purple, ripening to with a mixture of Larix gmelinii and Picea jezoensis,
bright or dark red brown; old cones grey. Seed scales accompanied by Betula sp. and with Pinus pumila
(15–)20–40(–45), ovate oblong, slightly recurved or and Juniperus communis var. saxatilis common in
straight, convex, flattening out when opened, 5–10 the understorey. In mountain ranges on the mari-
× 4–8 mm at mid-cone; surface smooth, striated on time seaboard of NE Asia conditions for mixed co-
abaxial side, sometimes strongly reddish pubescent, niferous forest are more optimal and here L. gmelinii
usually weakly pubescent or glabrous; upper margin grows with Abies nephrolepis, A. holophylla, Picea
entire, rounded or truncate, or weakly to strong- obovata or P. jezoensis, and Betula ermanii, locally
ly emarginate and slightly or strongly recurved; also with Juniperus, Pinus pumila or Rhododendron
base narrowed or slightly pedicellate. Bracts broad sp. in the understorey.
ligulate-­lanceolate, length ½ × seed scales, included
except at base of cone, but visible with opened seed
Uses
scales; apex trilobate with longer cusp. Seeds ovoid,
slightly flattened, 2–3 × 1–2 mm, light brown; seed Dahurian larch is an extremely important tim-
wings ovate-oblong, 5–6 × 3 mm, bright orange- ber tree in the Russian Far East, where it provides
brown, shining. ­building logs for traditional log houses, railway
sleepers, fences, and gates, as well as timber for con- Larix gmelinii (Rupr.) Kuzen. var. japonica (­Maxim.
struction, ship building (in Japan and Sachalin), ex Regel) Pilg., in Engler & Prantl, Nat. Pflanzen-
and the pulp industry. Its variability also extends to fam., ed. 2, 13: 327. 1926. Larix dahurica Turcz. ex
the suitability as a plantation tree for forestry out- Trautv. var. japonica Maxim. ex Regel, Gartenfl.
side its natural range; provenances from Siberia are 20: 105. 1871; Larix gmelinii (Rupr.) Kuzen. subsp.
prone to ‘late’ frosts (which actually means: lack of a japonica (Maxim. ex Regel) E. Murray, Kalmia 12:
proper cold winter that lasts until spring definitively 21. 1982. Type: Japan: Hokkaido, Oshima Prov.,
arrives), while those of Japan and even Sakhalin are ­Hakodate, cultivated on temple grounds, C. J. Maxi-
less likely to be frost damaged. Inter-specific hybrids mowicz s.n. (holotype not located, isotype K).
have been produced in cultivation in Denmark
and China, but probably have not been planted far 485
Description
beyond the trial nurseries. Due to the risk of frost
damage this species and its varieties are rare in cul- This variety differs from var. gmelinii in the follow-
tivation in more temperate parts of the world. ing characters: Seed cones (when opened) wider
than long, 1.2–2.5 × 1.5–2.8 cm; seed scales more
4 varieties are recognized: numerous, 18–25, ovate oblong, with emarginate
and (strongly) recurved apex. Young shoots dark
Larix gmelinii (Rupr.) Kuzen. var. gmelinii. Abies purplish brown, densely pubescent at first. Leaves
gmelinii Rupr., Beitr. Pflanzenk. Russ. Reiches 2: 56. slightly shorter: 1.5–2.5(–3) cm.
1845. Type not designated.
Distribution
Larix dahurica Turcz. ex Trautv., Pl. Imag. Descr. Fl.
Russ. 3, 7: 48, t. 32. 1846, non Lawson (1836). Russian Far East: Kuril Islands, Sakhalin.
Larix gmelinii (Rupr.) Kuzen. var. genhensis S. Y. Li TDWG codes: 31 KUR SAK
& Adair, Sida 16 (1): 183. 1994; Larix gmelinii (Rupr.)
Kuzen. f. genhensis (S. Y. Li & Adair) L. K. Fu & Nan
Conservation
Li, Novon 7 (3): 262. 1997.
IUCN: LC
Description
Larix gmelinii (Rupr.) Kuzen. var. olgensis (A. Hen­
Seed cones small, 0.8–2(–2.5) cm long, 0.8–1.5 cm ry) Ostenf. & Syrach, Pflanzenareale 2: 62. 1930.
wide when opened; seed scales few (ca. 15–20), Larix olgensis A. Henry, Gard. Chron., ser. 3, 57:
glabrous or weakly pubescent; upper margin from 109, f. 31–32. 1915. Type: Russia: Russian Far East,
entire and rounded to emarginate and slightly re- Primoriye, Olga, [“St. Olga”], C. J. Maximowicz s.n.
curved. (holotype K).

Distribution Description
NE China: Manchuria, Nei Monggol [Inner Mongo- The main differences with var. gmelinii are: Seed
lia]; Mongolia; Russian Fed.: E. Siberia, Russian Far cones larger, 1.8–3.5(–4.5) × 1.5–3 cm; seed scales
East (including Kamchatka). more numerous (16–30), dark red brown, convex,
TDWG codes: 30 BRY CTA IRK YAK 31 36 CHI-NM opening at 45° from the cone rachis. Young shoots
CHM 37 MON (densely) orange brown pubescent, but sometimes
nearly glabrous.
Conservation
Distribution
IUCN: LC
NE China: Jilin, E Liaoning; North Korea; Russian
Far East: Sikhote Alin Range.
TDWG codes: 31 PRM 36 CHM-JL CHM-LN 38 KOR-
NK
 Conservation Description
IUCN: LC Trees to 15–20(–23) m tall, d.b.h. to 0.5–0.8 m; trunk
monopodial, straight or bend; bark on trunk scaly,
Larix gmelinii (Rupr.) Kuzen. var. principis- with broad, shallow fissures and thick ridges, grey.
rupprechtii (Mayr) Pilg., in Engler & Prantl, Nat. Branches long, spreading horizontally, towards the
Pflanzenfam., ed. 2, 13: 327. 1926. Larix principis- ends assurgent, finally long pendulous; branches
rupprechtii Mayr, Fremdl. Wald-Parkbäume: 309. of second order long, slender, extremely pendu-
1906; Larix gmelinii (Rupr.) Kuzen. subsp. principis- lous; crown broad conical or pyramidal, in old trees
rupprechtii (Mayr) E. Murray, Kalmia 12: 21. 1982. domed. Branchlets long, slender, reddish brown or
486 Type not designated. Fig. 158 orange brown, becoming grey, yellowish or light
brown pubescent or glabrous, glabrous in the third
Larix principis-rupprechtii Mayr var. pendula year; short shoots stout, ringed with reflexed rem-
D. S. Zhang & Y. M. Chen, J. Beijing Forest. Univ. 10 nants of perular scales, 3–8 mm long. Vegetative
(2): 113. 1988; Larix gmelinii (Rupr.) Kuzen. f. pen- buds ovoid or conical, 2 × 1.5 mm, resinous; bud
dula (D. S. Zhang & Y. M. Chen) L. K. Fu & Nan Li, scales triangular, red brown. Leaveson short shoots
Novon 7 (3): 261. 1997. spirally, densely set in false whorls, 30–50 or more,
spreading radially, (1–)1.5–3.5(–4.5) cm long, 1–1.5
mm wide, narrowly linear, soft, flexible, more or less
Description
flat above, carinate below, obtuse-acutish at apex;
This variety differs from the others mainly by its stomata in a few interrupted lines above, in two nar-
large female cones (2–4 × 2–2.5 cm), which have row bands below; leaf colour green, shiny, stomatal
numerous seed scales (25–45) and resemble those bands greenish white, leaves yellow in autumn.
of L. decidua. The first year’s shoots are yellowish or Pollen cones terminal on short shoots, peduncu-
orange brown and usually glabrous, but sometimes late, pendant, often numerous, 1–2 cm long, yellow,
slightly brown pubescent. with red rachis, perular scales reddish brown. Seed
cones terminal on short shoots, erect from pendu-
lous branches; peduncles 1–2 cm, curved, scaly, with
Distribution
leaves; cones cylindrical, with obtuse apex, (3–)5–
North Central China: Hebei, Henan, N. Shanxi. 8(–11) cm long, 2.5–3.2 cm wide with opened scales;
TDWG codes: 36 CHN-HB CHN-SX CHS-HE colour (immature) purplish, with reddish purple
bracts completely covering the seed scales, ripen-
ing to dark brown, with purple brown, light fringed
Conservation
bracts. Seed scales more or less orbicular, convex or
IUCN: LC flat, often undulate, 10–15 × 10–15 mm; surface stri-
ated, with yellowish brown, short pubescence per-
sisting only on lower part; upper margin entire at
Larix griffithii Hook. f., Himal. J. 2: 44. 1854. first, soon erose, usually undulate and slightly re-
[validating descr. in vol. 1: 255] curved; base narrowed, often lacerate. Bracts broad
or narrow lanceolate, cusp elongated, recurved, edg-
es lacerate, 2–3 cm long, 5–8 mm wide, exserted and
Etymology
strongly or slightly reflexed. Seeds ovoid-cuneate,
This species has been named after William Griffith 5 × 3 mm, yellowish brown; seed wings ovate oblong,
(1810–1845), who when Director of the Botanic Gar- 6–10(–12) × 4–6 mm, purplish brown.
den of Calcutta amassed a large herbarium now held
at Kew.
Taxonomic notes
The two similar names Larix griffithii and L. griffit-
Vernacular names
hiana have been used more or less interchangeably
Sikkim larch; Xizang hongshan (Chinese) in the literature on conifers, in fact I used the lat-
ter in my book Pinaceae (Farjon, 1990). However,
the former (1854) has priority over the latter (1855), Larix griffithii Hook. f. var. griffithii. Type: Bhutan:
the presumed basionym Abies griffithiana Lindl. & [“Bootan”, locality not stated], W. Griffith 4989
Gordon (1850) being a nomen nudum, and must be (lectotype C). Fig. 159
applied to this species following the International
Code of Botanical Nomenclature. Abies griffithiana Lindl. & Gordon, J. Hort. Soc.
London 5: 214. 1850, nom. nud.
Larix griffithiana hort. ex Carrière, Traité Gén. Con-
Distribution
if.: 278. 1855.
E. Himalaya; China: NW Yunnan, SE Xizang [Ti- Larix kongboensis R. R. Mill, Novon 9 (1): 79. 1999.
bet].
TDWG codes: 36 CHC-YN CHT 40 EHM-AP EHM- 487
Description
BH EHM-DJ EHM-SI NEP
Shoots yellowish or light brown pubescent; leaves
(1–)1.5–3.5(–3.8) cm long, 1–1.2 mm wide. Bracts of
Ecology
seed cones strongly reflexed.
Larix griffithii occurs in the cloud belt of the
E Himalayas, at elevations between (1800–)2400–
Taxonomic notes
4000(–4100) m a.s.l., on rocky moraines or various
lithosols. The climate is moist to wet (summer mon- The smaller cones of Larix kongboensis (3–5 cm long)
soon), with annual precipitation exceeding 2000 are cited as an important character state separating
mm. It occurs in pure forests up to the tree line, at them from the larger cones of L. griffithii (Grimshaw
lower elevations it is often mixed with Abies specta- & Bayton, 2009: 438–439), but the cones of the latter
bilis, A. densa, Pinus wallichiana, Picea spinulosa, show great variation in length and we do not know
Tsuga dumosa and Juniperus sp. Betula utilis and if these smaller sizes (as well as the shorter dwarf
various large species of Rhododendron are the most shoots) are not due to slow growth at high altitudes
common broad-leaved trees associated with it. on the dry side of the Himalayas. More substantial
evidence is needed for the separation as a distinct
species of this small population in SE Xizang [Ti-
Uses
bet]. Rushforth (2009) cited its separation as a spe-
Sikkim larch is of minor economic importance as a cies, together with his recognition at that rank of
timber tree due to its occurrence in remote valleys Picea linzhiensis, in support of a third new conifer,
and on high slopes. It was introduced to Britain in Abies fordei Rushforth, all endemic to the Yarlung
the 19th century but was not very successful and re- Zangbo river drainage.
mains restricted to a few arboreta and other large
gardens with collections of exotic trees, usually in
Distribution
countries or regions with a mild climate and rare
occasions of frost. The main problem seems to be E Himalayas from E Nepal, Sikkim and Bhutan to
early flushing of leaves in regions with erratic warm Arunachal Pradesh (NE India), in the Chumbi Val-
spells in winter, which then get damaged by ‘late’ ley it reaches into Xizang [Tibet], China.
frosts. It would thus be expected to perfom better TDWG codes: 36 CHT 40 EHM-AP EHM-BH EHM-
in countries with a more continental, but not ex- DJ EHM-SI NEP
treme winter cold climate. The name Larix griffithi-
ana Carrière is still often applied to this species, but
Conservation
this is a later and therefore superfluous name for the
same species (see Taxonomic notes). IUCN: LC

2 varieties are recognized:

 Larix griffithii Hook. f. var. speciosa (W. C. Cheng
Vernacular names
& Y. W. Law) Farjon, World Checkl. Bibl. Conif.:
139. 1998. Larix speciosa W. C. Cheng & Y. W. Law, Japanese larch; Karamatsu (Japanese)
Acta Phytotax. Sin. 13 (4): 84. 1975.
Description
Description
Trees to 30–35(–40) m tall, d.b.h. to 1–1.5 m; trunk
Long shoots darker than in var. griffithii, glabrous; monopodial; bark on trunk scaly and fissured, dark
short shoots thicker; leaves 2.5–4.5 cm long, 1–1.5 reddish or purplish brown, with grey plates. Branches
mm wide. Bracts of seed cones narrower and less long, spreading horizontally, upper ones ascending;
488 strongly reflexed. branches of second order relatively short, slender,
drooping but not pendulous; crown broad pyrami-
dal, dense, or more open and irregular, conical and
Taxonomic notes
narrower in forest stands. Branchlets slender, flexi-
In Flora of China 4: 34 (1999) this variety is treated as ble, not pendulous, variable in colour, from yellowish
a distinct species. The distinctive characters appear to orange brown or purplish brown, bloomed with
to be all of a gradual, continuous nature, whereas grey, shining, glabrous or slightly pubescent; short
between species we must expect more discontinu- shoots cylindrical, with rings of perular scale bases,
ous morphological differences. Both varieties, ac- 4–10 mm long. Terminal buds conical, 4–5 mm long,
cording to Chinese accounts, occur on the Tibetan slightly resinous; bud scales triangular ovate, with
side of the Himalayan watershed. Unfortunately, the erose margins, dark red brown. Leaves on long shoots
disputed border area between India and China, as to 6 cm long and 2 mm wide; on short shoots spirally,
well as extreme N Myanmar [Burma], have long not in false whorls of 20–35, spreading radially, (2–)3–4
been visited by botanists making collections, so we cm long, 1 mm wide, linear, widest above the mid-
do not know whether this variety occurs in valleys dle, in cross section diamond shaped; apex obtuse;
draining to the south. stomata in several lines above, in two narrow bands
separated by a midrib below; leaf colour greyish or
bluish green, bright yellow in autumn. Pollen cones
Distribution
terminal on short shoots, often clustered on short
China: NW Yunnan, SE Xizang (Tibetan Himalaya). branches, 5–10 mm long, yellow. Seed cones termi-
TDWG codes: 36 CHC-YN CHT nal on short shoots, erect from drooping branches;
peduncles short, curved; cones broad ovoid, with
truncate apex, (1.5–)2–3(–3.5) cm long, 1.5–2(–2.5)
Conservation
cm wide with opened scales; colour (immature) vio-
IUCN: NT let, the bracts with a green midrib, maturing to yel-
lowish or orange brown, dark, dull brown when old.
Seed scales 30–40, sub-orbicular, concavo-convex,
Larix kaempferi (Lamb.) Carrière, Fl. Serres Jard. 10–13 × 10–13 mm at mid-cone, opening wide when
Eur. (Ghent) 11: 97. 1856. Pinus kaempferi Lamb., ripe, striated on both surfaces, glabrous; upper mar-
Descr. Pinus 2: [Pref.] v. 1824. Type: Illustration: gin entire, undulate or emarginate, usually strongly
Kaempfer, Delin. Pl. Japon.: t. 218 [ms.] in Bibl. recurved; base narrowed. Bracts ligulate, ending in
Sloan., Min. 139, Cat. No. 2914 xxvi G (Brit. Mus., a short cusp, half the length of seed scales, included,
Bloomsbury) (lectotype). Fig. 160 but visible in lower part of cone when seed scales are
opened. Seeds ovoid cuneate, slightly flattened, 4 ×
3 mm, brownish white, mottled with red; seed wings
Etymology
ovate oblong, 8 × 4 mm, reddish yellow with brown.
This species was named after Engelbert Kaempfer
(1651–1716), a German botanist and physician, and
Distribution
one of the earliest Europeans to describe and illus-
trate Japanese plants. Japan: central Honshu.
TDWG codes: 38 JAP-HN
 Larix laricina (Du Roi) K. Koch, Dendrol. 2 (2):
Ecology
263. 1873. Pinus laricina Du Roi, Diss. Inaug. Obs.
Larix kaempferi is a species of mesic sites, occurring Bot.: 49. 1771. Type not designated.
from the hills to high in the mountains (500 m to
2300 m a.s.l.), on the south face of Fuji san it reaches Larix alaskensis W. F. Wight, Smithsonian Misc.
2900 m. Unlike the other NE Asiatic larches it oc- Collect. 1 (50): 174. 1908; Larix laricina (Du Roi)
cupies better soils, often of recent volcanic origin, K. Koch var. alaskensis (W. F. Wight) Raup, Sargen-
and is never found on peat. The climate is cold, with tia 6: 105. 1947.
snowy winters and abundant rain in cool summers.
It is commonly found in association with other coni-
Etymology
fers, e.g. Pinus densiflora, Picea jezoensis subsp. hon- 489
doensis, Tsuga diversifolia, Abies homolepis at lower The species epithet means ‘similar to larix’ and was
elevations, and A. veitchii at higher elevations, but it given under the genus name Pinus, referring to Pi-
is clearly a sub-climax species. Several broad-leaved nus larix L.
tree genera are present at the lower elevations, e.g.
Quercus, Fagus and Betula. Pure ‘scrub stands’ may
Vernacular names
occur at the upper limit of trees.
Tamarack, Eastern larch, American larch
Conservation
Description
IUCN: LC
Trees to 30–35 m tall, d.b.h. to 0.8–1 m; trunk mo-
nopodial; bark on trunk very scaly, reddish brown
Uses
with small grey plates. Branches moderately long
Japanese larch is an important timber tree in Ja- or short, slender, spreading horizontally or curved
pan and in Europe (Scotland), where it has been downward; branches of second order relatively
introduced in 1834. The wood is similar to that of short, slender, drooping or pendulous; crown (nar-
European larch and is used for construction, rail- rowly) conical, broader in the south, narrower in
way sleepers, pit props and the pulp industry. It is the north. Branchlets slender, flexible, drooping or
also a frequently planted amenity tree in parks and lower ones pendulous, orange brown or purplish,
large gardens and a limited number of cultivars are pruinose, later blackish grey, glabrous or slightly pu-
known. In Scotland, a spontaneous hybrid occurred bescent at first; short shoots cylindrical, 3–10 mm
around 1900 between L. kaempferi and L. decidua long. Terminal buds ovoid, 3 × 2 mm, resinous; bud
which was named Larix ×eurolepis Henry (but is scales triangular obtuse, smooth, with ciliate mar-
correctly named Larix ×marschlinsii Coaz based gins, dark brown. Leaves on short shoots in false
on an earlier crossing event) and shows marked F1 whorls of (12–)15–20(–25), (1.5–)2–3(–3.5) cm long,
hybrid vigour or heterosis. Its seed cones resemble 0.5–1 mm wide, narrowly linear, diamond shaped
those of L. kaempferi with recurved scale apices, but in cross section, keeled, obtuse to acutish at apex;
are larger. This fast growing hybrid became much stomata none or a few faint lines above, 2 narrow
favoured by foresters and has been propagated and bands below; leaf colour light green or bluish green,
planted widely in many parts of Europe, often in- later dark green, stomatal bands greenish white,
volving back-crosses with either parents. Despite leaves turn bright yellow in autumn. Pollen cones
this greater production of timber per ha/year of the terminal on short shoots, small, yellow when ripe.
hybrid, Japanese larch remains an important planta- Seed cones terminal on short shoots, more or less
tion tree for timber on poorer soils, where neither erect; peduncles short, slender, curved; cones ovoid-
the hybrid not the other parent do so well and where globose, widening with opened scales, 1–2 cm long,
much of Europe’s plantation forestry is situated (the 1–2 cm wide with opened scales; colour (immature)
better soils being occupied by agriculture mostly for light red or greenish, maturing to orange brown
food crops). with a purplish tinge, old cones grey brown. Seed
 scales 10–20, orbicular or sub-orbicular, convex, from sapling trees. In the past, the northern tribes
8–10 × 7–9 mm at mid-cone; abaxial surface stri- of Native Americans used the roots of Tamarack to
ated, glabrous; upper margin serrulate or nearly en- sew birch bark canoes and the wood supplied the
tire, incurved; base narrowed. Bracts ligulate, with shafts for arrows. Its use as an amenity tree outside
tiny cusp, ca. ¼ length of seed scales, included, only its natural range is limited due to damage from
lower bracts visible. Seeds triangular ovoid, 3 × 2 ‘late’ frosts; however there should be some southern
mm, yellowish brown; seed wings ovate-oblong, 5–8 provenances that are more suitable.
× 3–4 mm, yellowish, translucent.

Larix lyallii Parl., Conif. Nov.: 3. Jan 1863. [&

Distribution
490 J. Bot. 1: 35. 1863]. Type: USA: Washington, North
N North America: from Newfoundland and Massa- Cascades, [“East side of Cascade Mts. latit. 49°”],
chusetts to Yukon and British Columbia, disjunct in D. Lyall s.n. (holotype K). Fig. 161
interior Alaska.
TDWG codes: 70 ASK NWT 71 ABT BRC MAN SAS
Etymology
72 LAB NBR NFL-NE 74 ILL MIN WIS 75 CNT INI
MAI MAS NWY OHI PEN VER This species was named after David Lyall (1817–1895)
who collected the type specimen.
Ecology
Vernacular names
This is mostly a species of the lowland boreal and
subarctic forests across Canada, it is less common Alpine larch, Subalpine larch
in medium high mountains. Its altitudinal range is
from 1 m to 1220 m a.s.l., but in British Columbia
Description
and Alaska it does not occur above 520 m. Tamarack
will grow on a variety of acid soils, but is found most Trees to 20–25 m tall, d.b.h. to 0.5–0.8 m; trunk
commonly on peaty soils in swamps and muskegs. monopodial; bark scaly and fissured below in old
The climate in its vast range is likewise varied, rang- trees. Branches moderately long, slender or mas-
ing from cool, moist maritime on the Atlantic coast, sive, ascending to nearly erect, or more horizontal;
to extremely dry, cold continental in the interior. It branches of second order short, flexible, not pen-
occurs locally in pure stands (maritime), but else- dulous; crown broad or narrow, conical, irregular
where commonly with Picea mariana, P. glauca, and open in old trees. Branchlets rather stout, firm,
Abies balsamea, or Pinus banksiana; boreal broad- orange red, but young shoots densely covered with
leaved trees such as Populus tremuloides and P. bal- lanate, yellowish hairs, in second year almost gla-
samifera occur usually after disturbance, Betula may brous, grey, later blackish grey; short shoots ovoid-
be represented with tree and shrub species. The conical or barrel-shaped, with grey, pubescent rings
shrub layer is often well developed, with various eri- and apex, 5–10 mm long. Vegetative buds ovoid glo-
caceous species. bose, 3 × 2 mm, not resinous, yellowish pubescent;
bud scales obtuse triangular, brown. Leaves on short
shoots spirally, close, in false whorls of 25–35(–40),
Conservation
(1.5–)2–3.3 cm long, 0.6–1 mm wide, narrowly linear,
IUCN: LC curved or twisted, more or less rhombic in cross sec-
tion, keeled on both sides, obtuse or acute at apex;
amphistomatic, but more lines of stomata below;
Uses
leaf colour light green, later bluish green, yellow
Tamarack produces durable, dense wood that is in autumn. Pollen cones terminal on short shoots,
used for outdoor purposes such as posts, railway 10–15 mm long, yellow. Seed cones terminal on short
sleepers, log cabins and mine shaft timbers. Its main shoots, more or less erect; peduncles short, thick,
industrial use is for pulp wood feeding the manu- curved; cones ovoid-conical to cylindrical, obtuse
facture of paper, in particular transparent envelope or truncate at apex, 3.5–5 cm long, 2–2.5 cm wide
windows. In Alaska, dog sled runners are made with opened scales; colour (immature) ­yellowish
pubescent, almost hidden by purplish red bract Larix mastersiana Rehd. & E. H. Wilson, in Sar­
scales, maturing to light or pale brown with purplish gent, Pl. Wilson. 2: 19. 1914. Larix griffithii Hook.
bracts. Seed scales ovate oblong or sub-orbicular, re- f. var. mastersiana (Rehd. & E. H. Wilson) Silba,
curved when opened, 10–14 × 6–12 mm at mid-cone; Phytologia Mem. 7: 39. 1984. Type: China: Sichuan,
abaxial surface densely pubescent when young, later Qionglai Shan, Wolong reserve [“Kuan Hsien”],
glabrous, finely striated or smooth; upper margin E. H. Wilson 906 (holotype A).
entire, rounded, not incurved, base cuneate or nar-
rowed. Bracts broad ligulate-lanceolate, narrowing
Etymology
into a long, caducous cusp, length 1.5 × seed scales,
exserted, straight; cusps recurved. Seeds triangu- The epithet commemorates Maxwell T. Masters
lar or ovoid, flattened, 4 × 3 mm, light brown; seed (1833–1907), a physician and honorary botanist at 491
wings ovate oblong, 8–10 mm long, light brown or the Royal Botanic Gardens, Kew, who contributed
rose-brown, tinged with red. much to the study of conifers.

Distribution Vernacular names

Canada: SW Alberta, SE British Columbia; USA: Masters larch; chuan hong shan (Chinese)
N Idaho, W Montana, N Washington.
TDWG codes: 71 ABT BRC 73 IDA MNT WAS
Description
Trees to 20–25 m tall, d.b.h. to 0.8 m; trunk mono­
Ecology
podial, straight or curved; bark on trunk irregularly
Larix lyallii is a subalpine larch which occurs at or fissured and breaking into grey plates. Branches
near tree line, at elevations between 1220 m and long, the topmost ascending, the lower spreading
2440 m a.s.l. It grows usually on shallow, rocky horizontally or descending; branches of second
mountain soils, but occasionally on deeper, well order slender, pendulous, but less so than those of
drained soils if there are no competitive species. The L. potaninii; crown usually broad, domed in free
climate is cold, with short, cool summers and long, standing trees, or more conical in forest stands.
snowy winters. It may occur in pure stands or mixed Branchlets long, slender, pendulous, yellowish or
with e.g. Abies lasiocarpa, Pinus albicaulis, P. flexilis, reddish brown, becoming grey, glabrous, or very
Picea engelmannii, and Tsuga mertensiana, forming young shoots sparsely pubescent; short shoots cy-
small groves near the tree line or scattered, solitary, lindric, with rings of revolute scale bases, 3–15 mm
stunted trees, sometimes surrounded by very little long. Vegetative buds conical or ovoid, 2 × 1.5 mm,
vegetation taller than alpine meadows. resinous; bud scales triangular, light brown, shin-
ing. Leaves on on short shoots spirally, densely set
in false whorls of 20–40, (1.2–)2–3(–3.5) cm long,
Conservation
1 mm wide, narrowly linear, slightly wider near
IUCN: LC the obtuse to acutish apex, more or less rhombic in
cross section, keeled on both sides, most clearly near
base; stomata none or only a few faint lines above,
Uses
in 2 narrow bands below; leaf colour bright green,
Due to its usually scattered occurence at high alti- yellow in autumn. Pollen cones terminal on short
tude Subalpine larch has no commercial value as a shoots, pedunculate, erect or pendant, 10–15 mm
timber tree. Its wood properties are similar to oth- long, yellow. Seed cones terminal on short shoots,
er larches. It establishes as a pioneer in avalanche more or less erect from pendulous branches; pe-
chutes and can minimize the destructive impact of duncles curved, 5 mm long; cones ovoid-cylindrical,
snow avalanches better than most other conifers. curved or straight; apex obtuse, 2.5–4.5 cm long, 1.5–
In horticulture, it is very susceptible to ‘late’ frosts 2.5 cm wide with opened scales; colour (immature)
and therefore seldom planted except in regions with greenish, with orange yellow bracts covering seed
long, consistently cold winters. scales, maturing to light brown, with dark brown
bracts, old cones dull, dark brown, with blackish
 bracts. Seed scales 30–40, obcordate-orbicular, con- It has been over-exploited in its natural range, espe-
vex, 6–10 × 7–12 mm at mid-cone; surface smooth cially in more accessible localities. Outside China,
or slightly striated, puberulent when young, later where it is used in afforestation, this species is not in
glabrous; upper margin entire, slightly emarginated, cultivation except for a few specimens in living tree
base very short pedicellate or cuneate. Bracts broad, collections (arboreta).
lanceolate, exposed part triangulate, reflexed, with
lacerate margins, cuspidate, 2–2.3 cm long, exserted.
Seeds ovoid-cuneate, 3 × 2 mm, yellowish brown Larix occidentalis Nutt., N. Amer. Sylva 3: 143, t.
or light brown; seed wings obovate, 6–8 × 4–5 mm, 120. 1849. Type not designated.
light yellowish brown.
492
Etymology
Distribution
The species epithet means ‘from the west’.
China, W Sichuan (Motian Ling, Min River drain-
age, Jiajin Shan).
Vernacular names
TDWG codes: 36 CHC-SC
Western larch
Ecology
Description
Larix mastersiana is a high mountain species of rare
occurrence, its altitudinal range is between 2000 m Trees to 45–55(–65) m tall, d.b.h. to 1.5–2(–2.2) m;
and 3500 m a.s.l. It grows in podzolic mountain soils, trunk monopodial, straight; bark on trunk scaly,
usually on steep slopes with good drainage. The cli- rough, deeply fissured below, grey brown, old bark
mate is cold-temperate and moist. It has not been plates grey. Branches relatively short, slender, spread-
reported from climax forest with other conifers, ing horizontally or curved down below, assurgent to-
but seems to be a tree thriving in secondary growth wards the top; branches of second order short, stout,
after forest clearing, as inferred from photographs the smallest slender, drooping to pendulous; crown
by E. H. Wilson taken at the beginning of the 20th narrowly conical, especially so in old trees, dense or
century [collection of the Arnold Arboretum, Mass., open. Branchlets slender, flexible, the longest becom-
USA and a 4-volume photo-album entitled “Vegeta- ing pendant, (pale) orange brown or reddish brown,
tion of Western China” (Wilson, 1912), copy in the turning grey, glabrous or sparsely pubescent when
Herbarium of the Natural History Museum at Paris]. young; short shoots cylindrical, blackish grey, 4–15
mm long. Vegetative buds ovoid globose, 3 × 2 mm,
resinous; bud scales triangular, dark reddish brown.
Conservation
Leaves on long shoots to 6 cm long, on short shoots
Exploitation beyond sustainability has led to serious spirally, close, in false whorls of (15–)30–40(–45),
decline of this species in the more accessible parts of spreading radially, (1.5–)2–4(–4.5) cm long, 0.5–1
its limited natural range. It is now largely confined mm wide, narrowly linear, soft and flexible, in cross
to the steeper and higher localities where forest road section triangular, flattened, keeled below; apex ob-
building has not advanced. Recent Chinese policies tuse or acutish; hypostomatic, stomata in two narrow
to discontinue or at least restrict logging of the natu- bands separated by a midrib or keel below; leaf colour
ral forests of Sichuan and other western regions may, bright green, later dull green, turning bright yellow
if effective, halt or even reverse the decline. Afforesta- in autumn. Pollen cones terminal on short shoots,
tion using this species outside its natural range is not 8–15 mm long, yellow. Seed cones terminal on short
a true substitute for conservation of wild ­populations. shoots, more or less erect from pendulous branches;
IUCN: VU (A1c) peduncles 5–10 mm, curved; cones ovoid oblong or
conical, with obtuse apex, 2.5–4(–6) cm long, 2–2.5
(–3) cm wide with opened scales; colour (immature)
Uses
purplish red, with green bract scales, maturing to
A timber tree used for construction, pit props, rail- violet-brown or reddish brown, finally light brown
way sleepers and furniture; the bark yields tannins. or grey-brown. Seed scales 35–45, cuneate-orbicular,
emarginate at apex, or sometimes rounded, 8–14 × Larix potaninii Batalin, Trudy Imp. S.-Peterburgsk.
08–12 mm at mid-cone; surface smooth, abaxial side Bot. Sada 13: 385. 1894. Pl. 20
finely striated, glabrous; upper margin entire, often
deeply emarginated, base narrowed or short pedicel-
Etymology
late. Bracts ligulate, tapering to an elongated cusp,
length ca. 2 × seed scales, exserted with recurved This species was named after the Russian botanist
cusps. Seeds ovoid-triangular, 3–4 × 2.5 mm, light Grigorii N. Potanin (1835–1920), who travelled and
brown, with dark spots; seed wings obliquely ovate, collected in western China in the 1880’s.
6–9 × 4–5 mm, pale yellowish brown.
Vernacular names
493
Distribution
Chinese larch; hongshan, hungsha (Chinese)
Canada: British Columbia; Pacific NW of USA.
TDWG codes: 71 BRC 73 IDA MNT ORE WAS
Description
Trees to 40–50 m tall, d.b.h. to 1–1.5 m; trunk mo-
Ecology
nopodial, straight or curved; bark on trunk rough
Larix occidentalis occurs in the mountains, at eleva- and scaly, dark brown in fissures, plates grey. Branch-
tions between 600 m and 2100 m a.s.l., usually on es long, ascending or spreading, drooping at the
grey brown, well drained podzolic mountain soils, ends; branches of second order long, slender, pen-
which are moderately acid. The climate is cold, with dulous; crown (broad) conical or domed. Branch-
cool summers and moist winters, the annual pre- lets slender, firm, later long, flexible and pendulous,
cipitation ranges from 450 mm to 875 mm, much of deep reddish brown or yellowish orange, with grey
it falls as snow. It may occur in pure stands; in an grooves, later grey, glabrous, or with some scattered
initial stage after disturbance (e.g. fire) Pinus con- hairs in the grooves; short shoots small, cylindri-
torta var. latifolia can become dominant, followed cal, 3–5 × 3–8 mm. Vegetative buds ovoid globose,
by P. ponderosa in certain areas; later in the succes- 3 × 2.5 mm, resinous; bud scales obtuse triangular,
sion P. monticola, Pseudotsuga menziesii, Abies gran- with erose margins, dark red brown. Leaves on short
dis and A. lasiocarpa, finally Thuja plicata and Tsuga shoots spirally, dense, in false whorls of 20–40, 1.2–
heterophylla take their place. Picea engelmannii and 3.5 cm long, ca. 1 mm wide, narrowly linear, wid-
Tsuga mertensiana occur mainly above Larix occi- est above the middle, soft, more or less triangular in
dentalis and may be associated with L. lyallii. cross section, flattened, keeled below, obtuse acutish
at apex; stomata in several faint lines above, in two
narrow bands below; leaf colour bright green, turn-
Conservation
ing yellow in autumn. Pollen cones terminal on short
IUCN: LC shoots, ca. 10 mm long, yellow. Seed cones terminal
on short shoots, erect from pendulous branches; pe-
duncles short, curved; cones cylindrical or elliptical,
Uses
obtuse at apex, (2.5–)3–5(–9) cm long, 1.5–2.5(–3.5)
Western larch is an important timber tree. It can cm wide with opened scales; colour (immature) vio-
grow to great size with straight boles and grows let, with purple bracts, ripening to dark brown, with
rapidly in height though it takes longer to increase purplish black bracts. Seed scales 35–65(–80), sub-
in girth. The wood is durable, hard and strong and orbicular, 9–13 × 9–12 mm at mid-cone; surface fine-
used for long poles, railroad sleepers, mine timbers, ly striate or smooth, glabrous; upper margin entire
fine veneer, and pulpwood for the paper industry. or erose, round or truncate, not incurved; base very
The resin from the wood has useful water soluble short pedicellate. Bracts broad ligulate-lanceolate;
properties for a variety of industrial products espe- apex acute or mucronate-cuspidate, 1.2–2.2 cm ×
cially applied in ink, paint and offset lithographic 4–5 mm, exserted, straight, dark purple or blackish,
printing. The use of this species in amenity planting with lighter midrib. Seeds triangular-ovoid, slightly
is limited, although some provenances should grow flattened, 3 × 2.5 mm, pale brown, with dark spots;
well in cooler climates.
494

5
6

Pl ate 2 0. Larix potaninii. 1. Habit of tree. 2. Branchlet with foliage. 3. Branch with seed cones (var.
potaninii). 4. Seed cone (var. macrocarpa). 5. Seeds. 6. Seed scale with bract.
seed wings ovate oblong, 6–8 × 4–5 mm, brown,
Distribution
tinged with purple.
China: S Gansu, S Shaanxi, W Sichuan, NW Yun-
nan, E Xizang [Tibet].
Distribution
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHN-
China: S Gansu, S Shaanxi, W Sichuan, NW Yun- SA CHT
nan, E & S Xizang [Tibet]; Nepal.
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHN-
Conservation
SA CHT 40 NEP
IUCN: LC
495
Ecology
Larix potaninii Batalin var. chinensis (Beissn.)
Larix potaninii and its varieties are high mountain L. K. Fu & Nan Li, Novon 7 (3): 262. 1997. Larix
larches occurring between 2350 m and 4300 m al- chinensis Beissn., Mitt. Deutsch. Dendrol. Ges. 1896
titude a.s.l. The soils are acidic mountain podzols. (5): 68. 1896, non Mill. (1768). Type: China: Shanxi,
The climate is cold, the precipitation varies from 800 [northern Shanxi, “Kuon-tou-san” (mountain)],
mm to 2000 mm annually according to location. J. Giraldi 6 (lectotype K).
At high elevations it grows often pure, with under-
growth of Juniperus squamata, elsewhere it is usu-
Description
ally mixed with Abies spp., Picea spp., Tsuga dumosa
or T. chinensis, Cephalotaxus spp. and Taxus spp. Short shoots 3–4 mm diam., densely yellow pubes-
cent. Seed cones 2.5–5 cm long, 1.5–2.8 cm wide
when opened; seed scales striate abaxially.
Uses
Chinese larch is an important timber tree in the
Distribution
western mountains of China. Its wood is durable
and can be used for construction purposes, mining China, S Shaanxi (Taibai Shan).
props and railway sleepers, as well as milled for ve- TDWG codes: 36 CHN-SA
neer and pulped for the paper industry. It is rarely
used in forestry plantations outside China and for
Conservation
amenity planting. It is in cultivation only as speci-
men trees in some arboreta and pineta; most trees in IUCN: LC
Europe and the USA derived from seed collected by
Ernest Wilson and other plant collectors early in the Larix potaninii Batalin var. himalaica (W. C. Cheng
20th century and often have not survived. There are & L. K. Fu) Farjon & Silba, Phytologia 68: 37. 1990.
numerous recent collections planted, however these Larix himalaica W. C. Cheng & L. K. Fu, Acta
are still confined to arboreta. This species should be Phytotax. Sin. 13 (4): 84. 1975. Type: China, Xizang
planted more often in suitable regions (climate) as [Tibet], north side of Xomolungma [Mt. Everest],
it has large, attractive cones with conspicuous up- Chinese collector No. 80A (holotype PE).
turned bracts.
Description
4 varieties are recognized:
Seed cones max. 6.5 cm long; bracts mucronate cus-
Larix potaninii Batalin var. potaninii. Type not pidate. Young shoots yellowish orange.
designated.
Taxonomic notes
Description
In Flora of China 4: 34 (1999) this variety has been
Seed cones 3.5–5.5 cm long, 1.5–3 cm wide when treated as a distinct species. The morphological dif-
scales are opened; seed scales 35–65 in number. ferences appear to be of a quantitave and more or
 less gradual nature and it shares the typical upright
Vernacular names
bracts of L. potaninii.
Siberian larch; Listvennitsa sibirskaya (Russian);
Xinjiang luoye song (Chinese)
Distribution
China: S Xizang [Tibet]; Nepal (Langtang Khola).
Description
TDWG codes: 36 CHT 40 NEP
Trees to 30–40 m tall, d.b.h. to 1–1.5; trunk mono­
podial, straight or curved; bark on trunk light
Conservation
brown, rough and scaly. Branches spreading hori-
496 This variety apparently occupies a limited area as it zontally, near the top ascending; branches of second
is known with certainty only from some valleys in order slender, long, drooping or pendulous; crown
the Mt. Everest area, both on the Chinese and Nepa- broad conical, often irregular. Branchlets slender,
lese side of the border. In Xizang [Tibet] it occurs in flexible, pale yellowish, lustrous, usually glabrous
Chomolungma National Park. or with minute pubescence in grooves; short shoots
IUCN: VU (D2) conical-globose or short cylindrical, 4–10 mm long.
Vegetative buds ovoid, 3 × 2 mm, not resinous; bud
Larix potaninii Batalin var. macrocarpa Y. W. Law, scales ovate, more or less pubescent, dark greyish
Acta Phytotax. Sin. 13 (4): 84. 1975. Type: China: brown. Leaves on short shoots spirally, close, in
Yunnan, locality not known, Chinese collector No. false whorls of 20–40, (2–)2.5–4(–5) cm long, 0.5–1
9347 (holotype PE). mm wide, narrowly linear, soft, flexible, flattened,
faintly keeled below, obtuse or acutish at apex; sto-
mata in a few lines above, in two narrow bands be-
Description
low; leaf colour (light) green, turning bright yellow
Seed cones 5–8(–9) × 2.5–3.5 cm; seed scales 50–80 in autumn. Pollen cones terminal on short shoots,
in number. 5–10 × 5–6 mm, pale yellow. Seed cones terminal
on short shoots, more or less erect from pendulous
branches; peduncles thick, curved, 5–10 mm long;
Distribution
cones ovoid or subglobose, (2.5–)3–4.5(–5.5) cm
China: SW Sichuan, NW Yunnan. long, (1.8–)2.5–3.5(–4) cm wide with opened scales;
TDWG codes: 36 CHC-SC CHC-YN colour (immature) reddish or rose-green, ripening
to (light) red-brown, old cones grey-brown. Seed
scales 25–40, sub-orbicular to ovate, strongly con-
Conservation
vex or less so in transitional forms, 12–20 × 12–18
IUCN: LC mm at mid-cone; surface smooth or slightly stri-
ated, in the typical form densely ferruginous pubes-
cent, the exposed part of scales becoming glabrous
Larix sibirica Ledeb., Fl. Altaica 4: 204. 1833. Larix with age; upper margin entire, rounded, truncate or
decidua Mill. var. sibirica (Ledeb.) Regel, Gartenfl. slightly emarginate, usually incurved; base rounded
20: 101. 1871; Larix decidua Mill. subsp. sibirica or cuneate. Bracts ligulate-linear, length ¼-⅓ × seed
(Ledeb.) Domin, Acta Bot. Bohem. 10: 6. 1931. Type: scales, included. Seeds ovoid-cuneate, flattened, 5 ×
Russia: Siberia, Altai Mts., C. F. von Ledebour et al. 3.5 mm, (light) brown with dark spots; seed wings
s.n. (holotype LE). oblique-ovate, 8–16 × 4–8 mm, lustrous red-brown
or orange-brown.
Pinus larix L. var. russica Endl., Syn. Conif.: 134.
1847; Larix russica (Endl.) Sabine ex Trautv., Trudy
Taxonomic notes
Imp. S.-Peterburgsk. Bot. Sada 9 (1): 212. 1884.
The correct name for this species is Larix sibirica
Ledeb. (1833) as it is the earliest binomial published
Etymology
under Larix for this species. In my book Pinaceae
The species epithet refers to its origin in Siberia. (Farjon, 1990) I incorrectly used L. russica (Endl.)
Sabine ex Trautv., based on Pinus larix L. var. rus- peat or on mountains above the steppe (Altai Mts.),
sica Endl. (1847) as the accepted name. This name but more common it is mixed with Pinus sylvestris,
was also used in Dallimore & Jackson’s well known P. sibirica, Picea obovata, Abies sibirica and broad-
Handbook of Coniferae and Ginkgoaceae (e.g. edition leaved trees such as Betula pendula and Populus spp.
4, 1966) and in other compilations of conifers. This
species was long considered a mere form or variety
Conservation
of the European larch, L. decidua, first described as
Pinus larix by Linnaeus (1753). When raised to spe- IUCN: LC
cies rank, it is compulsory to take up the earliest
species epithet published for it, which was given by
Uses
Ledebour in his Flora Altaica (op. cit.). 497
Siberian larch is an important timber tree in Russia.
It is logged from natural coniferous forests as well
Distribution
as from plantations, which are established in Rus-
Russia: from the White Sea to Lake Baikal in Siberia; sia outside its natural range as well as in Finland,
China: Xinjiang (Altai Mts., E Tien Shan); Mongolia Poland and Sweden. The wood is durable and used
(Altai Mts.). in construction, traditionally for log houses in Si-
TDWG codes: 14 RUN RUE 30 ALT IRK KRA TVA beria for which the wood is roughly hewn to shape,
WSB 36 CHX 37 MON but untreated. It has been widely used for railroad
sleepers e.g. on the famous Trans Siberian Railroad.
Larch wood is also milled for construction timber
Ecology
and veneer and pulped for the paper industry. Sibe-
Larix sibirica is common in the lowland taiga of rian larch is in cultivation as an amenity tree, but it is
W Siberia, where it forms the northern limit of trees vulnerable to ‘late’ frosts in climatic zones with mild,
alternately with Picea obovata and with Pinus sylves- fluctuating winter temperatures. For this reason it is
tris. It also occurs in the mountains (from 500 m to not used in western Europe but successfully planted
2400 m a.s.l.). It grows on a great variety of soils, in central and eastern Europe, in Scandinavia, and
from peat bogs to well drained, sandy or rocky soils, as far afield as Iceland. A few cultivars are known,
where it has its optimum. The climate is very cold but with limited distinction from habit forms that
(min. temp. –55 °C), continental or subarctic, dry, also occur in nature.
with very long winters. There are pure stands on
 Lepidothamnus Phil., Linnaea 30: 730. 1861. Type: Lepidothamnus fonkii Phil.
(Podocarpaceae).

Greek: lepis, lepidos = scale; thamnos = bush, shrub. Lepidothamnus fonkii Phil., Linnaea 30: 731. 1861.
Dacrydium fonkii (Phil.) Benth. & Hook. f., Gen. Pl.
3 (1): 433. 1880. Type: Chile: Aisén, Archipelago de
Description
los Chonos, F. Fonk 263 (holotype SGO). Fig. 162
Creeping or erect (dwarf) shrubs or trees, dioecious
or sometimes monoecious, evergreen. Resin canals
Etymology
498 in leaves absent (unique in family). Bark on trees
thin, with minute lenticels, becoming scaly and flak- This species was named after Francis Fonk, who col-
ing. Branching irregular, ascending in trees. Leaves lected the type specimen in 1857.
spirally arranged or opposite-decussate in juvenile
plants, dimorphic, with linear, spreading juvenile
Vernacular names
leaves usually gradually giving way to subulate, de-
current leaves and finally appressed, keeled, ovate- Chilean rimu [rimu is the Mauri (New Zealand)
rhombic and gibbous scale leaves. Stomata on all name for Dacrydium cupressinum, so this one is a
sides (leaves amphistomatic). Pollen cones terminal trifle far-fetched but nevertheless recorded in books
or sometimes lateral and axillary on scale-leaved on (cultivated) conifers].
branchlets, solitary, short cylindrical; microsporo-
phylls spirally inserted, triangular, with two globose
Description
pollen sacs at base containing bisaccate pollen. Seed
cones terminal, solitary, very small, consisting of Dioecious or occasionally monoecious dwarf shrubs
3–5 bracts slightly larger but similar to scale leaves, to 50 cm tall, creeping or sometimes more or less
becoming distinct by colouring yellow or reddish, erect. Stems mostly decumbent, with foliage branch-
swelling to become succulent or not, 1–2 bracts fer- es ascending to erect. Juvenile leaves restricted to
tile with an erect, axillary ovule. Seeds 1 per cone, seedlings, decurrent, often distichous, acicular, 4–6
maturing in the second year, ovoid, not compressed, mm long, bilaterally flattened, slightly curved, acute.
ripening to lustrous purplish brown or black and Adult leaves spirally arranged, on leading branches
surrounded at base by a membranous, greenish scale-like, appressed, ovate-oblong, keeled towards
sheath formed by the epimatium. apex, 4–7 mm long, 2–3.5 mm wide, on ultimate
branchlets scale-like, imbricate, 1.5–3 mm long, 1–1.5
3 species. mm wide, strongly keeled to gibbous towards the
distal part of branchlets, with membranous mar-
gins; apex slightly incurved, obtuse. Stomata on
Distribution
adaxial side of juvenile leaves numerous from base
S Argentina; S Chile; New Zealand. to apex, only a few short lines on abaxial side near
base, on adult leaves conspicuous on abaxial side,
scattered but mostly absent on keel and apex. Pol-
Key to the species of Lepidothamnus
len cones solitary, terminal, sessile, 5–6 × 1.5–2 mm;
1a. Erect shrubs or small trees to 15 m tall microsporophylls 12–16, triangular, with acute apex
L. intermedius and two reddish basal pollen sacs. Seed cones soli-
1b. Prostrate, creeping or sometimes erect dwarf tary, terminal, consisting of 3–5 leaf-like bracts with
shrubs to 50 cm tall 2 a narrow base, distal 1–2 bracts fertile. Bracts may
2a. Adult scale leaves on ultimate branchlets 1–1.5 or may not fuse and swell to form a succulent, red
× 1 mm, slightly keeled. Native in New Zea- receptacle. Seeds at base enclosed by a membranous
land L. laxifolius epimatium, 4–5 mm long, ovoid-oblong with an api-
2b. Adult scale leaves on ultimate branchlets 1.5–3 culate apex, maturing to dark brown or black.
× 1–1.5 mm, strongly keeled to gibbous. Native
in Chile and Argentina L. fonkii
 Distribution Uses
S Argentina: Chubut, Santa Cruz; S Chile: Aisén, There are no economic uses of this species. In cul-
Los Lagos, Magallanes. tivation it is only represented in a few botanic gar-
TDWG codes: 85 AGS-CB AGS-SC CLS-AI CLS-LL dens and perhaps an occasional private collection. It
CLS-MG should make an interesting and quite hardy ground
covering, evergreen conifer shrub in wet areas
around ponds or lakes, but, judging from its natu-
Ecology
ral habitat, evidently prefers low pH water and sub-
This dwarf conifer occurs in Sphagnum bogs and strate and a paucity of nutrients.
moorlands; its stems are usually hidden in moss or 499
other vegetation and only upright foliage branches
protrude. It can form quite extensive low thickets Lepidothamnus intermedius (Kirk) Quinn, Austral.
that exclude all or most other plants, rooting on the J. Bot. 30 (3): 316. 1982. Dacrydium intermedium
decumbent stems. Its altitudinal range varies with Kirk, Trans. New Zealand Inst. 10: 386. 1878. Type:
latitude as it occurs roughly between 40° S and 55° S, New Zealand: South Island, Westland, Hokitika,
between 900 m in the north to near sea level in the T. Kirk 806 (syntype K, lectotype not designated).
south. In the north it is growing in bogs in forested
areas with Pilgerodendron uviferum, Fitzroya cupres- Dacrydium intermedium Kirk var. gracilis Kirk,
soides and Nothofagus antarctica. The latter ‘antarc- Trans. Proc. New Zealand Inst. 17: 224. 1885. Type:
tic beech’ is virtually the only tree at its southern New Zealand: Stewart Island, Ruggedy, T. Kirk 1072
limit, in between Pilgerodendron uviferum also oc- (lectotype K, designated here).
curs frequently. Sedges (Carex spp.) and other plants
characteristic of these bogs are also common.
Etymology
The species epithet refers to intermediacy of char-
Conservation
acters compared to other species of Lepidothamnus.
This species is threatened mainly in the northern
part of its range, where it is associated with Fitzroya
Vernacular names
cupressoides and/or Pilgerodendron uviferum. As
a result of extensive exploitation of these tree spe- Yellow silver pine
cies, habitats have been modified (usually drained
at minimum, in many areas converted to pasture).
Description
However, it is unlikely that similar habitat degrada-
tion has occurred where these conifer tree species are Shrubs or small trees to 15 m tall; trunk to 60 cm
either absent (Fitzroya cupressoides) or of a smaller d.b.h., often multi-stemmed. Bark on larger trunks
stature no longer of interest for their timber. This is exfoliating in irregular, thin flakes; freshly exposed
the case in the southern part of the range (Magal- bark purplish brown, older patches turning dull
lanes) of L. fonkii and it can be assumed that the spe- brown and finally grey-white, densely pitted with
cies is there fairly safe. What is less well known is its brown dots. Branches ascending, forming a broad,
overall abundance in this botanically little explored, rounded and more or less open crown. Juvenile
remote region and what proportion of the total area leaves on seedlings and young plants, distinct from
of occupancy (AOO) these occurences represent. adult leaves (but intermediate forms occur), ac-
A comprehensive geographical survey of this species icular, spreading at near right angles to shoot, 6–8
(not an easy task given the vastness and remoteness (–10) mm long, more or less four-ranked, straight
of the Chilean southern archipelago) is wanting. or slightly curved, slightly bilaterally flattened, ta-
IUCN: VU (B1+2c) pering to an acute apex. Intermediate leaves shorter,
3–5 mm long, triangular in cross-section, tapering
to an acute or apiculate apex. Adult leaves gradually
appearing above these, decurrent at base, spreading
towards the free apex on most vegetative branchlets,
 appressed on ultimate branchlets with seed cones or and the ‘grass trees’ Dracophyllum latifolium or
on dense sun-exposed foliage of old trees, 1.2–2 × 1 D. pyramidale on Great Barrier Island.
mm (appressed leaves 1 × 1 mm), keeled abaxially,
obtuse. Stomata on all sides of juvenile and adult
Conservation
leaves, in two indistinct bands separated by the keel
on adult leaves. Pollen cones terminal, sessile, cy- IUCN: LC
lindrical, 5–6 × 2 mm; microsporophylls triangular
with a constricted, more or less free apex, with two
Uses
basal red pollen sacs. Seed cones terminal, consist-
ing of a short axis with 4–6 leaf-like bracts which The wood of the Yellow silver pine was formerly
500 turn yellow, the distal one fertile, not swelling. Seeds used for railway sleepers and telegraph poles due to
surrounded at base by a drying epimatium with its strength and durability. It is reddish yellow and
erose margin, erect, ovoid, 4–5 × 2.5 mm, constrict- contains much resin in the sapwood, which prob-
ed at apex, green ripening to lustrous purplish black. ably renders it highly flammable. As with most other
native trees in New Zealand, commercial exploita-
tion has ceased after most of the lowland forests
Taxonomic notes
were destroyed and is now prohibited. This species
Thomas Kirk described and named Dacrydium is very rare in horticultural cultivation, it may be
intermedium var. gracilis from Stewart Island. At present in a few botanic gardens in New Zealand
K there are three small specimens on one sheet and abroad; in colder climates it would likely only
alledgedly collected by him and numbered 1071– grow under glass, but in milder ones it may be more
1073. The collecting date of No. 1071 is given as De- or less hardy depending on provenance.
cember 1883 but Kirk visited the island only twice,
in January 1882 and again in January 1884. Nos. 1072
and 1073 are undated as to collection time but were Lepidothamnus laxifolius (Hook. f.) Quinn, Aus-
“communicated” in May 1884. These can have been tral. J. Bot. 30 (3): 316. 1982. Dacrydium laxifolium
collected by Kirk in January 1884 and then sent to Hook. f., London J. Bot. 4: 143. 1845. Type: New
J. D. Hooker at Kew. Brian Molloy selected No. 1072 Zealand: North Island, Wellington, Tongariro N. P.,
as the lectotype of Kirk’s variety on this sheet in Sep- Tongariro Volcano, near the summit, J. C. Bidwill
tember 1991 and that specimen is here designated 5; W. Colenso 60 (syntypes K, lectotype not desig-
as the lectotype of D. intermedium var. gracilis Kirk. nated). Pl. 21
Both of us have concluded upon examination that
this variety is not taxonomically different from the
Etymology
species, Lepido­thamnus intermedius (Kirk) Quinn.
The species epithet means ‘with soft (lax) leaves’, but
the leaves are rather stiff. It may instead describe the
Distribution
foliage shoots.
New Zealand: North Island, South Island, Stewart
Island.
Vernacular names
TDWG codes: 51 NZN NZS
Pygmy pine
Ecology
Description
Lepidothamnus intermedius is most common on
Stewart Island, where it forms the main shrub or Low, usually prostrate dioecious or monoecious
tree in swamp forests. On the west coast of South dwarf shrubs forming mats or trailing through veg-
Island and in North Island it is more scattered and etation. Stems slender, creeping, rarely sub-erect, to
grows in boggy terrain on slopes and sometimes on 1 m long, occasionally branching. Foliage branches
hill summits and ridges to ca. 900 m a.s.l. The spe- very slender and flexuous, 1–2 mm thick, spread-
cies is accompanied by tall shrubs or tussock ­grasses ing or ascending, sparse or more crowded. Leaves of
two types, juvenile and adult (intermediates occur),
 6

501

1
5

Pl ate 2 1. Lepidothamnus laxifolius. 1. Habit of shrub. 2. Branch with foliage and seed cones. 3. Branchlet
with juvenile leaves. 4. Juvenile leaves. 5. Adult leaves. 6. Pollen cone. 7. Seed cone and seed.
 often mixed on a single leading branch but adult
Ecology
leaves distal from juvenile leaves. Juvenile leaves
alternate, subulate to linear, stiff and spreading at Lepidothamnus laxifolius is one of the smallest coni-
more or less right angles to shoot, 3–8 mm long, fers. It occurs in moorlands, peat bogs, and tussock
keeled abaxially, flat or slightly concave adaxially, grass slopes in the mountains, usually between 750
obtuse. Transitional leaves shorter but similar, 1.5–3 m and 1200 a.s.l., on Stewart Island down to sea level.
mm long, spreading at less than 90°. Adult leaves It can form extensive mats creeping over rocks and
spirally arranged, imbricate, appressed, oblong- long runners trailing through the vegetation. It is as-
ovate, 1–1.5 × 1 mm, slightly keeled, obtuse. Stomata sociated with tussock grass (Chionochloa rubra) and
on juvenile leaves numerous on adaxial side, in two with Halocarpus bidwillii, H. biformis, Podo­carpus
502 sparse bands on abaxial (keeled) side; scattered on nivalis, Hebe (Veronica s. l.), Olearia, Dracophyllum,
adult leaves. Pollen cones solitary, terminal, sessile, Pseudopanax, Gleichenia, Phormium, and other
5–7 × 2 mm; microsporophylls broadly triangular shrubby plants and ferns.
with acute-apiculate apex and two basal red pollen
sacs. Seed cones solitary, terminal, consisting of a
Conservation
short axis with 3 slightly spreading bracts that fuse
and swell to a globose, succulent (but sometimes dry IUCN: LC
and not swollen) receptacle and turn orange-red.
Seeds enclosed at base by an epimatium, 4–5 mm
Uses
long, ovoid-oblong, with a small, curved apiculus,
ripening brown or purplish brown with longitudinal Pygmy pine has no commercial value, but would be
lighter striations. an interesting and probably excellent dwarf shrub
for rock gardens. It is currently only present in a few
botanic gardens and perhaps an occasional private
Distribution
collection. With plants only 7–8 cm long known to
New Zealand: North Island, South Island, Stewart have borne ripe seeds, it may well be the smallest
Island. conifer species known.
TDWG codes: 51 NZN NZS
Libocedrus Endl., Syn. Conif.: 42. 1847. Type: Libocedrus plumosa (D. Don) Sarg.
[Dacrydium plumosum D. Don] (Cupressaceae).

Stegocedrus Doweld, Novosti Sist. Vyssh. Rast. 33: 42. (Gadek & Quinn, 1993; Brunsfeld et al., 1994) and
2001. Type: Stegocedrus austrocaledonica (Brongn. & on matK sequences (Gadek et al., 2000) it was found
Gris) Doweld [Libocedrus austrocaledonica Brongn. that there is no close relationship between Libo­
& Gris]. cedrus s. str. and Calocedrus, the northern hemi-
sphere genus that was long included. Phylogenetic
Greek: libos = tear, drop (referring to resin exuda- analysis based on morphology as well as on “com-
tions); Cedrus is the classical name for (true) cedars. bined data” (Gadek et al., 2000) confirmed this, so 503
there is strong support for at least the separation of
these two. Phylogenetic relationships are much clos-
Description
er with the southern hemisphere taxa, but how close
Shrubs or trees to 35 m, evergreen, monoecious, and with which taxa has not been unambiguously
multistemmed or monopodial. Bark scaly, exfolia- demonstrated. Here the data have so far produced
ting in longitudinal strips or plates, reddish brown or conflicting results in phylogenetic analysis. Molecu-
brown. Branches spreading or ascending, forming a lar data have placed Pilgerodendron nested within
pyramidal, conical or bushy crown. Foliage branch- Libocedrus (Gadek et al., 2000) but morphology did
es frondose and plagiotropic, sometimes more ir- not (Hill & Brodribb, 1999; Farjon, 2005a).
regularly disposed, spreading or ascending. Leaves
scale-like, decussate, decurrent, imbricate, strongly
Key to the species of Libocedrus
dimorphic on flattened (pen)ultimate branchlets,
facials smaller than laterals or nearly equal in size; Leaves should be examined on ultimate branchlets
facials rhombic, 1–5 mm long; laterals divergent, of full grown, not young plants.
2–7 mm long, conduplicate, falcate, both eglandu-
lar; margins entire; apex free or appressed, obtuse to 1a. Large trees. Pollen cones with 8–14 microsporo­
acute; stomata predominantly in conspicuous bands phylls. Native to New Zealand 2
on underside of branchlets. Pollen cones terminal, 1b. Shrubs or small trees. Pollen cones with 16–24
solitary, 2.5–10 × 2–3.5 mm; microsporophylls 8–24, microsporophylls. Native to New Caledonia 3
decussate, peltate, bearing 4(–6) abaxial pollen sacs. 2a. Ultimate branchlets more or less quadrangular,
Seed cones terminal on flattened or quadrangu- with facial leaves only slightly smaller than lat-
lar branchlets, subtended by 4–5 transitional leaf eral leaves. Free part of the bract a third of the
pairs, 8–18 mm long; bract-scale complexes forming size of the seed cone scale and not exceeding
the cone in 2 decussate pairs; upper fertile pair of beyond the cone scale L. bidwillii
scales spreading at maturity, with a long subapical 2b. Ultimate branchlets remain flattened, with fa-
bract tip; lower pair similar but smaller; columella a cials less than half the size of laterals. Free part
small spike. Seeds 1–4 per cone, with 2 very unequal of the bract half the size of the seed cone scale
wings. Seedlings with 2 cotyledons. and exceeding beyond the cone scale
L. plumosa
5 species. 3a. Leaves on ultimate branchlets of nearly equal
size. Lower pair of seed cone scales only slightly
smaller than upper pair L. chevalieri
Distribution
3b. Leaves on ultimate branchlets very unequal
New Zealand, New Caledonia. in size. Lower pair of seed cone scales much
smaller than upper pair 4
4a. Apex of facial leaves not reaching the next fa-
Taxonomic notes
cial leaf above, obtuse. Bract of lower, smaller
Phylogenetic relationships among genera in Cupres- seed cone scales twice as long as these
saceae based on DNA sequence data have only more L. austrocaledonica
recently been analysed. Based on rbcL sequences
 4b. Apex of facial leaves reaching the next facial ­ eveloping within one growing season to become
d
leaf above, apiculate. Bract of lower, smaller thin woody, 10–12 mm long. Bract-scale complexes
seed cone scales as long as these L. yateensis 6–8 × 3–5 mm, slightly rugose, spreading free parts of
the bracts 5–7 × 1 mm on laterals and 8–10 × 1.5 mm
on facials. Columella a small spike ca. 1 mm long.
Libocedrus austrocaledonica Brongn. & Gris, Seeds 1–2, ovoid-oblong, slightly flattened, with an
Bull. Soc. Bot. France 18: 140. 1871. Stegocedrus acute to bifid apex, 5–6 × 2.5 mm, light brown, with
austrocaledonica (Brongn. & Gris) Doweld, Novosti 2 opposite, thin membranous wings, the smaller a
Sist. Vyssh. Rast. 33: 42. 2001. Type: New Caledonia: narrow strip less than 1 mm wide, the larger oval-
Grande Terre, Province Sud, Mont Humboldt, oblong, 6–8 × 2.5–3 mm, yellowish brown.
504 B. Balansa 2503 (holotype P).
Distribution
Etymology
New Caledonia: Province Sud, Province Nord (Mt.
The species epithet refers to its (predominant) oc- Paéoua).
currence in the southern part of New Caledonia. TDWG codes: 60 NWC

Vernacular names Ecology

No common names have been recorded for this spe- In low, shrubby rain forest on exposed ridges of
cies. the higher mountains at altitudes between 750 m
and 1400 m a.s.l., with Falcatifolium taxoides, Neo­
callitropsis pancheri, Podocarpus spp. Prumnopitys
Description
ferruginoides and here and there emergent Arau-
Shrubs or small trees 2–4(–6) m tall, multistemmed caria humboldtensis; also with various angiosperms
or sometimes monopodial, diam. ca. 10 cm. Bark e.g. Myrtaceae, Trimeniaceae, and Winteraceae, with
rough and scaly, reddish brown, exfoliating in longi- a well developed understorey of mainly ferns and
tudinal thin fibrous strips or plates. Branches sparse, abundant epiphytic mosses and lichens. The soils are
slender, spreading, foliage branches more numerous, mostly skeletal, strongly humic and very acid; this
spreading nearly horizontally, forming a narrowly species seems to avoid ultramafic rock types. Rain-
conical crown in young trees but irregular in shrubs. fall above 1000 m altitude usually exceeds 3500 mm
Foliage frondose, ultimate branchlets opposite, of and the vegetation remains under a blanket of cloud
equal length but towards end of foliage branches most of the time. Temperatures in these tropical
gradually shortening, entirely covered with leaves, mountains are therefore substantially cooler than at
persistent. Leaves decussate, on leading shoots long lower altitudes.
decurrent, on lateral (ultimate) branchlets adnate at
base, imbricate, strongly dimorphic, facials small,
Conservation
visible part 1–2 mm, rhombic-apiculate, keeled, ap-
pressed, partly covered at base by much larger 3–6 This species is restricted to high mountains in south-
(–7) × 1.5–3 mm, divergent, bilaterally flattened, fal- ern New Caledonia and one mountain in the north.
cate laterals with scarious margins; amphistomatic, The populations are small; regeneration seems to be
stomata on facials near base, on laterals much re- absent in the northern locality. Wildfires and min-
duced on upperside, in a conspicuous band on un- ing operations (there is a causal relationship be-
derside, upperside lustrous olive green or slightly tween the two through access roads to remote areas)
glaucous, underside with glaucous white stomatal are the major causes of threat. Substantial popula-
band. Pollen cones terminal, solitary, (ovoid-)ob- tions are protected in the Parq National de la Rivière
long, 5–8 × 2–2.5 mm; microsporophylls decussate, Bleue; protection of the population on Mt. Paéoua in
16–24, peltate with acute apex, weakly keeled, with Province Nord is urgent as it occurs near a mining
entire margins, bearing 4 small abaxial yellow pollen concession.
sacs. Seed cones terminal on flattened branchlets, IUCN: NT
 to ovoid, 2.5–5 mm; microsporophylls decussate,
Uses
8–10(–14), peltate, with entire margins, bearing
No uses have been recorded of this species and, 4 abaxial yellow pollen sacs. Seed cones terminal on
apart from a few plants in botanic gardens, it is not branchlets with monomorphic leaves of equal size;
known to be in cultivation. upper pair of leaves developing within one growing
season, becoming thin woody, together 8–12 mm
long. Bract-scale complexes 7–10 mm long, slightly
Libocedrus bidwillii Hook. f., Handb. New Zealand rugose, recurved in upper half above abaxially ex-
Fl. 1: 257. 1864 Type: New Zealand: South Island, serting bract tip, subtended by the lower, smaller
Marlborough, Richmond Range, [“Nelson Mts.”], (3–4 mm) less modified pair. Columella present, in
J. C. Bidwill [ex herb. W. J. Hooker] 126 (holotype K). mature cones a small spike 1–2 mm long. Seeds 2–4, 505
Fig. 163, 164 ovoid, flattened, with an acute apex, 2–3 mm long,
brown, with a whitish hilum and 2 opposite, thin
membranous wings; smaller wing a narrow strip
Etymology
less than 1 mm wide; larger wing irregularly oval-
This species was named after John C. Bidwill (1815– oblong, 4–5 × 2–3 mm, yellowish brown.
1853), who was an early botanical collector in New
Zealand and has sent many specimens to William
Distribution
Hooker, the first Director at the Royal Botanic Gar-
dens, Kew. New Zealand: North Island, South Island.
TDWG codes: 51 NZN NZS
Vernacular names
Ecology
Cedar; pahautea (Maori)
Libocedrus bidwillii is a co-dominant or canopy
emergent tree in montane-subalpine evergreen
Description
rainforests of the mixed angiosperm-conifer class,
Trees to 25(–28) m tall, monopodial, d.b.h. to ca. growing together with the podocarps Dacrydium
1–1.5 m. Bark thin, scaly, greyish brown, exfoliating cupressinum (at lower altitudes), Halocarpus bi-
in longitudinal thin strips. Branches long, spreading formis, Phyllocladus trichomanoides var. alpinus, and
or ascending, foliage branches numerous, arranged Podocarpus cunninghamii. Frequent angiosperm
in dense tufts above each other, forming a pyrami- trees are Metrosideros umbellata, Nothofagus solan-
dal crown in young trees, conical or irregular with a dri, Quintinia acutifolia and Weinmannia racemosa.
clear bole in forest stands. Foliage in flattened sprays Libocedrus bidwillii is a good example of a long-lived
in young trees, in old trees more irregular and as- conifer (max. 800–1000 years) dependent for regen-
cending, ultimate branchlets subopposite to alter- eration on stand-replacing episodal disturbances.
nate, 5–40 mm long, entirely covered with leaves, The altitudinal range is from 250 m to 1200 m a.s.l.
changing with age of plant from flattened to more The soil in these forests has a high organic content
or less quadrangular, persistent. Leaves decussate, and is usually saturated with water. In many places
on lateral (ultimate) branchlets short decurrent, at higher altitudes the forest is confined to drainage
imbricate, dimorphic in young trees, with facials systems, surrounded by peaty moorland dominated
small, rhombic, 1.5–2 × 1 mm, apiculate to acute, ap- by tussock-forming sedges. The climate is super­
pressed, partly covered at base by larger 2–4(–6) × humid, with high rainfall and frequent fog, sum-
1.5–2.5 mm, divergent, bilaterally flattened, slightly mers are short and cool.
curved laterals with entire margins and free apices,
leaves on older trees smaller and nearly monomor-
Conservation
phic; amphistomatic, stomata on facials near base,
on laterals much reduced on upperside, in a short, This species is not threatened with extinction. Near-
conspicuous band on underside, upperside dull ly all remaining natural ‘old-growth’ stands are now
(dark) green, underside with whitish green stomatal protected.
band. Pollen cones terminal, solitary, subglobose IUCN: LC
 equally ­distributed on both faces; leaf colour (light)
Uses
green, often with reddish brown in older leaves.
The reddish wood of this species is not very valu- Pollen cones terminal, solitary, cylindrical, 8–10 ×
able for timber as it splits easily. As an ornamental 2.5–3.5 mm; microsporophylls decussate, 16–24, pel-
tree it has been planted more often than Libocedrus tate with apiculate apex, weakly keeled, with entire
plumosa, the lowland species of New Zealand, but it margins, bearing 4(–6) small abaxial yellow pollen
remains restricted to arboreta and other tree collec- sacs. Seed cones terminal on branchlets, developing
tions in large gardens. within one growing season to become thin woody,
(10–)12–16 mm long. Bract-scale complexes slightly
different in size, with upper pair 10–14 × 5–7 mm
506 Libocedrus chevalieri J. T. Buchholz, Bull. Mus. and lower pair 10–12 × 3–4 mm, slightly rugose, ex-
Hist. Nat. (Paris), sér. 2, 21: 283. 1949. Stegocedrus cluding the spreading free parts of the bracts which
chevalieri (J. T. Buchholz) Doweld, Novosti Sist. are 6–8 × 1 mm on the larger pair and 5 × 1 mm on
Vyssh. Rast. 33: 42. 2001. Type: New Caledonia: the smaller pair. Columella a small spike ca. 1 mm
Grande Terre, Province Sud, Mont Humboldt, long. Seeds 1–2, ovoid-oblong, slightly flattened,
J. T. Buchholz 1567 (holotype ILL). with an acute apex, 5–6 × 2.5 mm, yellowish brown,
with 2 opposite, thin membranous wings; smaller
wing a narrow strip less than 1 mm wide; larger wing
Etymology
oval-oblong, 8–10 × 3–4 mm, often curved, yellow-
This species was named after L. Chevalier, who col- ish brown.
lected plants in New Caledonia in the 1940s and
1950s.
Distribution
New Caledonia: Province Sud (Mt. Humboldt, Mt.
Vernacular names
Kouakoué), Province Nord (Poindimié).
No common names have been recorded for this spe- TDWG codes: 60 NWC
cies.
Ecology
Description
On slopes near the summits of some of the high-
(Spreading) shrubs or small trees 2–5 m tall, mul- est peaks in usually steep terrain covered by a 2–5
tistemmed or sometimes monopodial, diam. ca. m tall ‘maquis’ vegetation, in the contact zone be-
10 cm. Bark rough and scaly, brown, exfoliating in tween schists and serpentines. The altitudinal range
thin irregular strips or plates. Branches numerous, is from 650 m to 1620 m a.s.l. These mountain sum-
ascending, forming a dense, bushy, often round- mits receive high levels of precipitation.
ed crown. Foliage more or less frondose, forming
dense sprays, ultimate branchlets alternate to sub-
Conservation
opposite, of nearly equal length but towards end of
foliage branches gradually shortening, 2–5 cm long, This rare species is known from only three very
rhombic in cross-section, slightly flattened, 3–4 mm small populations on isolated mountain summits,
wide, entirely covered with leaves, persistent. Leaves where growth is slow and regeneration poor. There
decussate, on leading shoots broad decurrent, es- is risk of wildfires even though two of the three pop-
pecially laterals apically recurved or spreading, on ulations are within floristic reserves. There is a likeli-
ultimate branchlets imbricate, weakly dimorphic hood of mining in the unprotected area.
to nearly monomorphic; facials and laterals nearly IUCN: CR [B2ab(iii)]
equal in size, 2.5–5 × 2–2.5 mm; facials triangular-
rhombic, appressed, keeled near apex, apiculate,
Uses
partly covered at base by bilaterally flattened, api-
cally more or less recurved laterals with acute-acu- No uses are recorded of this species; it is grown in
minate apex; amphistomatic, stomata on facials near only a few botanic gardens, as young plants in re-
the base, on laterals more conspicuous, more or less search greenhouses.
Libocedrus plumosa (D. Don) Sarg., Silva N. Amer. rugose abaxial surface. Columella present in mature
10: 134. 1896. Dacrydium plumosum D. Don, in cones, a small conical spike 2–3 mm long. Seeds 2–4,
Lambert, Descr. Pinus, ed. 3, App.: --. 1832. Type: ovoid, flattened, with an acute apex, 3–5 mm long,
New Zealand: locality unknown, A. Cunningham brown, with a whitish hilum and 2 opposite, thin
330 (neotype K). Fig. 165 membranous wings; smaller wing a narrow strip
less than 1 mm wide; larger wing irregularly oval-
oblong, 6–8 × 3–4.5 mm, yellowish brown.
Etymology
The species epithet (Latin plumosus) means ‘feath-
Distribution
ery’ and refers to the appearance of the foliage.
New Zealand: North Island, in South Island restrict- 507
ed to the Nelson district.
Vernacular names
TDWG codes: 51 NZN NZS
kawaka (Maori)
Ecology
Description
This species occurs in the lowland evergreen rain-
Trees to 30(–35) m tall, monopodial, d.b.h. to 2–3 m. forests of the mixed angiosperm-conifer class,
Bark thin, scaly, light brown to greyish brown, ex- where it is a canopy tree with other conifers, e.g.
foliating in longitudinal, ca. 10 cm wide, thin strips. Dacrycarpus dacrydioides, Dacrydium cupressinum,
Branches long, spreading or ascending, foliage Halocarpus kirkii, Manoao colensoi, Phyllocladus
branches numerous, spreading, arranged in dense trichomanoides, Podocarpus cunninghamii, P. to-
sprays above each other, forming a pyramidal crown tara, Prumnopitys ferruginea, P. taxifolia and in the
in young trees, conical, rounded or irregular with a far north of North Island Agathis australis. Undis-
clear bole in forest stands. Foliage in flattened sprays turbed forest of this type can have as many as eight
except in cone-bearing branches, ultimate branch- conifer genera (and species) on a single hectare, but
lets subopposite to alternate, 15–35 mm long, en- forest clearance as well as selective logging of ‘pines’
tirely covered with leaves, flattened, 2–6 mm wide, have drastically reduced these species-rich forests
persistent. Leaves decussate, on lateral (ultimate) especially in the lowlands. Various angiosperms
branchlets short decurrent, imbricate, dimorphic, are mixed in, e.g. Beilschmiedia tarairi, Dysoxylum
with facials small, rhombic, 1–2 × 1 mm, apiculate spectabile, and Leptospermum scoparium, but coni-
to acute, appressed, partly covered at base by larger fers (especially Agathis) can form groves with few
2–5(–6) × 1.5–2 mm, divergent, bilaterally flattened, angiosperms, forming a mozaic pattern rather than
slightly curved laterals with entire margins and an evenly mixed forest. Especially in gaps tree ferns
free apices, leaves on older trees smaller and nearly can become abundant. The altitudinal range is from
monomorphic; amphistomatic, stomata on facials near sea level to 600 m a.s.l. These forests receive
near base, on laterals in a small groove on upper- abundant rainfall throughout the year and tempera-
side, in a broad, conspicuous band of irregularly but tures are mild in winter and warm in summer.
densely arranged stomata on underside; upperside
green, underside with whitish green stomatal band.
Conservation
Pollen cones terminal, solitary, subglobose to ovoid,
3–5 mm; microsporophylls decussate, 8–12, peltate, Lowland forests have been greatly reduced since
with entire margins, bearing 4 abaxial yellow pollen ­European settlement in New Zealand began about
sacs. Seed cones terminal on branchlets with weakly two centuries ago. There is no quantitative record
dimorphic leaves of nearly equal size, developing available to indicate the reduction rate for this spe-
within one growing season to become thin woody, cies as a result of this wholesale removal of indig-
12–18 mm long. Bract-scale complexes 10–15 mm enous natural forest. Long term survival of this
long, with upper pair slightly recurved in distal half species in natural ecosystems requires large tracts
and with an obtuse to truncate apex, subtended by of unmanaged old growth forest, where natural cy-
the smaller (5–9 mm) acute pair, each with long ex- clic processes of disturbance and regeneration can
certed, reflexed and upcurved bract tips and finely span many centuries (see e.g. Ogden & Stewart in
 Enright & Hill, 1995). Such forests are still, despite Leaves decussate, especially laterals apically recurved
the creation of large reserves, under threat of being or spreading, on ultimate branchlets imbricate,
‘managed’ with selective logging one of the permit- strongly dimorphic; facials 1.5–2 mm long, rhombic,
ted uses. As with other long-lived conifers which act appressed, keeled near apex, apiculate, partly covered
as episodal pioneers, this form of management is in- at base by divergent, bilaterally flattened, falcate-lan-
compatible with their long-term existence. ceolate 2–5 × 1–2 mm laterals with entire to scarious
IUCN: NT margins and acute-acuminate apex; amphistomatic,
stomata on facials near base, on laterals reduced on
upperside, more conspicuous on underside in a de-
Uses
pressed band extending to apex; leaf colour (lustrous)
508 The wood of this species is dark reddish brown, fine- olive green, stomatal band glaucous white. Pollen
grained and often beautifully figured and therefore cones terminal, solitary, cylindrical, 5–8(–10) × 2–2.5
prized for furniture and wood panelling. Its rarity mm; microsporophylls decussate, 16–24, peltate with
and at least in recent times protection from exploi- apiculate apex, keeled, with entire margins, bearing 4
tation causes it to be of little economic value. As an small abaxial yellow pollen sacs. Seed cones terminal
ornamental tree it is uncommon, as it is less hardy on branchlets with leaves of nearly equal shape and
than its congener Libocedrus bidwillii, but it should size, developing within one growing season, becom-
do well in California, southern Europe, and areas ing thin woody. Bract-scale complexes much differ-
with a similar mild climate, provided there is no ent in size; upper pair 8–9 × 3–4 mm; lower, smaller
prolonged dry period. pair 4–6 × 2 mm; both slightly rugose, forming a cone
9–10 mm long excluding the erect free parts of the
bracts which are 5–8 × 1 mm on larger pair and 4–5
Libocedrus yateensis Guillaumin, Bull. Mus. × 1 mm on smaller pair. Columella a small spine less
Hist. Nat. (Paris), ser. 2, 21: 457. 1949. Stegocedrus than 1 mm long. Seeds 1–2, ovoid-oblong, slightly
yateensis (Guillaumin) Doweld, Novosti Sist. Vyssh. flattened, 5–6 × 2.5 mm, light brown, with 2 opposite,
Rast. 33: 42. 2001. Type: New Caledonia: Grande thin membranous wings; smaller wing a narrow strip
Terre, Province Sud, Rivière Bleue, J. Bernier s.n. less than 1 mm wide; larger wing oval-oblong, 6–7 ×
(lectotype P). 2.5–3 mm, often curved, yellowish brown.

Etymology Distribution
The species epithet indicates the locality, Yaté, near New Caledonia: Province Sud (Rivière Bleue-Yaté
where it was found when first described. River, Ouinné River), Province Nord (Povila).
TDWG codes: 60 NWC
Vernacular names
Ecology
No common names have been recorded for this spe-
cies. This species is restricted to riparian habitat along a
few rivers at low altitude, from 150 m to 600 m a.s.l.,
where both individual trees and small groves occur.
Description
It is one of only two species of Cupressa­ceae that
Shrubs or small trees 2–10(–12) m tall, multistemmed occurs in the lowland tropics; the other is Neocalli­
or monopodial, diam. ca. 10–30 cm. Bark rough and tropsis pancheri, which may occasionally grow with
scaly, reddish brown, exfoliating in thin longitudi- it but also occurs in the mountains. Libocedrus
nal strips or plates. Branches spreading to ascend- ­yateensis grows on sediments of river terraces or on
ing, forming a conical young tree but later a bushy, river banks, surrounded by various shrubs and ferns.
often rounded crown. Foliage more or less frondose,
forming dense sprays, ultimate branchlets alternate
Conservation
to subopposite, unequal in length and towards end of
foliage branches gradually shortening, flattened, ca. The populations of this species are small and each
4 mm wide, entirely covered with leaves, ­persistent. of them is vulnerable to fires. Most are protected in
the Parq Rivière Bleue de Yaté, but not those in the
Uses
vicinity of the Ouinnée River and a recently discov-
ered population in central New Caledonia at Roche No uses are recorded of this species; it is grown in
Ouaième. only a few botanic gardens, as young plants in re-
IUCN: EN [B2ab(iii)] search greenhouses.

509
 Manoao Molloy, New Zealand J. Bot. 33: 196. 1995. Type: Manoao colensoi (Hook.)
Molloy [Dacrydium colensoi Hook.] (Podocarpaceae).

Manoao is the Maori name for at least one other rib on both sides, straight or curved, stomata in two
podocarp (Halocarpus kirkii) as well as for this one, distinct bands mainly on adaxial side; apex acute.
between which the Maori presumably saw no dis- Branchlets with juvenile leaves may re-appear among
tinction. those with adult leaves. Intermediate phase leaves
often on ‘sucker’ shoots, 4–5 mm long, distichous,
bilaterally flattened. Adult leaves spirally arranged,
Description
510 imbricate and appressed, rhomboid in appearance,
See the species description. 1–1.5 × 1 mm, keeled abaxially; apex obtuse. Leaves
amphistomatic, stomata conspicuous, scattered. Pol-
len cones terminal, sessile, 4–6 mm long, 2–2.5 mm
Distribution
wide; microsporophylls (6–)8–12, rhombic to trian-
As for the species. gular, with minutely denticulate upper margins and
with two basal red pollen sacs. Seed cones terminal,
solitary or sometimes in pairs on distally curved
Manoao colensoi (Hook.) Molloy, New Zealand short branchlets, 3–4 mm long, consisting of 2–5(–6)
J. Bot. 33: 196. 1995. Dacrydium colensoi Hook., fertile bracts. Seeds 1–3(–5) per cone, crowded, mor-
Icon. Pl., n.s., 2: t. 548. 1843; Lagarostrobos colensoi phologically and topographically erect, ca. 3–4 × 2–3
(Hook.) Quinn, Austral. J. Bot. 30 (3): 317. 1982. mm, rounded in cross-section, notched ar apex, dark
Type: New Zealand: North Island, [“High hills near purple to black, basal half or more of seed enclosed
the eastern coast…”], W. Colenso 27 (lectotype BM). in a swollen, fleshy, yellowish green epimatium.
Fig. 166
Taxonomic notes
Etymology
Brian Molloy (op. cit.) segregated the New Zealand
The species epithet commemorates the missionary species (originally described by William Hooker as
William Colenso (1811–1899) who collected many Dacrydium colensoi) from Lagarostrobos, a genus
plants in New Zealand. erected by Chris Quinn (1982) to include this spe-
cies and L. franklinii from Tasmania. He chose a new
genus name, Manoao, the Maori name for the Sil-
Vernacular names
ver pine. Subsequent DNA-based investigations by
Silver pine Quinn and co-workers alledgedly refuted this seg-
regation, but with only two species (‘terminal taxa’)
represented in any cladistic analyses they will come
Description
out as sister taxa at their closest (as they do, see Con-
Evergreen, predominantly dioecious trees to 20 m ran et al., 2000) and at what rank you then recognize
tall; trunk to 0.7(–1) m d.b.h., often strongly tapered. them remains a matter of judgement and could in-
Bark on mature trees exfoliating in large scales, volve phyletic considerations (Stuessy, 2009). Mol-
leaving distinct wave and hammer marks, grey or loy has given quite a number of differences between
grey-brown. Crown of young trees broadly pyrami- the two, including numbers of chromosomes, and
dal, of old mature trees rounded, with large and together with their likely long (>85 million years)
steeply ascending main branches. ‘Sucker’ shoots separation across a widening ocean, we may as well
from horizontal underground stems prolific. Foli- recognize them as two separate genera. That these
age branchlets with adult leaves slender, 1.2–1.5 mm genera thereby become monotypic I do not consider
diam. including scale leaves, variable in length from a problem (e.g. Gregg’s Paradox: genus = species);
0.5–5 cm, spreading. Juvenile leaves on seedlings given the likely age of the separation event we can
and young plants distinct from adult leaves, acicular, postulate more than a single species in each genus,
6–10(–12) mm long, bifacially flattened with a mid- all but one now extinct.
 in New Zealand lowland rain forest, in South Island
Distribution
(Westland) it is often mixed with Nothofagus.
New Zealand: North Island, South Island, Stewart
Island.
Conservation
TDWG codes: 51 NZN NZS
IUCN: LC
Ecology
Uses
Manoao colensoi is a component of lowland conifer-
dominated rainforest occurring from near sea level The wood of Silver pine is dense and compact but
to ca, 950 m. It is shade tolerant and grows on well not heavy, strong and not shrinking when seasoned, 511
developed soils of volcanic or igneous origin with and taking a high polish. It is sometimes beautifully
good drainage. In places with high precipitation it figured or mottled and is then highly prized for cabi-
can occur outside forest cover, overtopping scrub. It net making and furniture. As native trees are now
is a very slow growing species which can maintain protected in New Zealand, its commercial use is
itself through all successional phases from forest dis- negligable. It is not known to be in cultivation out-
turbance to canopy tree in old growth. Its usual as- side a few botanic gardens.
sociates are other species in Podocarpaceae ­common
 Metasequoia Hu & W. C. Cheng, Bull. Fan Mem. Inst. Biol., ser. 2, 1 (2): 154. 1948
(nom. cons.). Type: Metasequoia glyptostroboides Hu & W. C. Cheng (Cupressaceae).

Greek meta- = changed; i.e. a changed or trans- proximally constricted and twisted, merging with
formed Sequoia, denoting its close relationship and alternating left and right trending decurrent bases,
similarity. lamina linear, curved or nearly straight, spreading
at nearly right angles to branchlet, 8–15 mm (–30
mm in young trees) × 1.5–2.5 mm, generally longest
Description
in middle section of branchlet, with parallel entire
512 See the species description. margins and a midrib, shortly tapering to obtuse or
mucronate apex. Leaves hypostomatic, stomata in
two broad bands of 4–8 lines divided by a narrow
Distribution
raised midrib, leaf colour light green or sometimes
As for the species. slightly glaucous green. Pollen cones numerous in
spike-like shoot systems, opposite, ovoid, 5–6(–10)
× 2.5–4 mm when full grown; microsporophylls
Metasequoia glyptostroboides Hu & W. C. Cheng, 15–20, proximally helically arranged, but irregularly
Bull. Fan Mem. Inst. Biol., ser. 2, 1 (2): 154. 1948. attached in mature cones, peltate, with 3 abaxial pol-
Type: China: Hubei, W Hubei, Modaoxi, [“E Sze­ len sacs near lower margin. Seed cones terminal on
chuan, Wanhsien, Mo-tao-hsi”], C. T. Hwa 2 (lecto- 2–5 cm long, sparsely (scale-)leaved shoots, solitary,
type NF). Fig. 167, 168 subglobose, barrel-shaped or fusiform when still
tightly closed, with nearly concentric grooves, glau-
Metasequoia glyptostroboides Hu & W. C. Cheng var. cous green, maturing in one year to 15–25(–30) ×
caespitosa Y. H. Long & Y. Wu, Bull. Bot. Res. North- 10–23 mm, parting the scales in the grooves, turning
East. Forest. Inst. 4 (1): 149. 1984. dull brown or grey-brown. Bract-scale complexes
16–24(–28), decussate, more or less peltate, dilated
into a transversely elliptic or broadly triangular disk
Etymology
with a transverse indentation, 5–8 mm long, 8–15
The species epithet refers to its likeness with Glyp- mm wide, 2–4 mm thick, distally rugose, proxi-
tostrobus. mally moderately lignified, striate, yellowish to dark
brown at base. Seeds numerous, more or less invert-
ed by displacement, 4–6 × 4–5 mm including wings,
Vernacular names
irregular, compressed, emarginate at apex, with two
Dawn Redwood; shui shan (Chinese) broad, membranous yellowish or light brown wings
encircling the seed.
Description
Distribution
Trees to 50 m tall (but most trees now under 35 m),
deciduous, monoecious; trunk monopodial, large Central China: Chongqing (Shizhu), Hubei (Li-
trees often buttressed, d.b.h. to 2.2 m. Bark becom- chuan, Zhonglu), Hunan (Longshan, Sangzhi).
ing fissured, exposing purplish brown inner bark, TDWG codes: 36 CHC-CQ CHC-HU CHC-SC CHS-
exfoliating in long, thin plates or strips. Branches HN
long, spreading to ascending, or foliage subpendu-
lous, forming a pyramidal crown in young trees,
Ecology
becoming broader and more open, with a rounded
top, in older trees. Foliage branchlets opposite or The ecology of Metasequoia in undisturbed valley
nearly so, mostly distichous, 5–15 cm long, 0.7–1 forests can only be reconstructed from palynologi-
mm thick including decurrent leaf bases, glabrous, cal data and from clues obtained from field observa-
terminating in small seasonal buds, deciduous. tions in the lower sections of some little side valleys
Leaves opposite on foliage branchlets, the free part of the Shuishaba Valley, where relatively undisturbed
stands occur on probably suboptimal sites. Remain- single tree, with little chance of natural generation
ing large trees in the valley proper are surrounded due to changes in land use by a growing agrarian
by cultivated fields (mainly rice) and both native population. Secondary woodland in some narrow
and introduced tree species, constituting in its most side valleys, where Metasequoia may occur along
natural state a secondary vegetation. Though some permanent streams, are under pressure of grazing
of the trees of Metasequoia have also been planted, it and cutting of firewood. The mature, large trees have
is likely that the largest and oldest trees in the valley all been declared protected but habitat protection is
are survivors. It is a riparian species that occupies overall inadequate (Bartholomew et al., 1983; Fu &
a habitat similar to that of Taxodium distichum; the Jin, 1992), which means that the survival of this very
remnant old trees may be the vestiges of an extensive interesting species in its natural habitat is not guar-
flood plain forest that existed before this valley was anteed. 513
transformed to agriculture only a few centuries ago. IUCN: CR (A1c, C2a)
Away from the valley floor the trees are restricted
to the moist bottoms of ravines and in contact with
Uses
seepage water. The valley forest would not have been
pure Metasequoia, but mixed with angiosperms, In the past, trees of this species must have been used
among which were very likely species of Acer, Cas- for construction timber locally. Evidence of branch
tanea, Populus, and Quercus, as well as Liquidambar cutting for firewood can be seen in the photograph
acalycina, Nyssa chinensis, Pterocarya hupehensis of one of the earliest discovered trees, a picture that
and other trees tolerant of periodic flooding. The has been widely published. Its use is now prohibited
soil is clay and sand derived from sandstone, with in the ‘Metasequoia area’ but the species has been
slightly acid to neutral pH and a strongly fluctuating widely planted, as an amenity or forest tree in Chi-
but not deep water table. The climate is character- na and as an ornamental in many other countries
ized by hot summers and cold winters. with temperate climates. This conifer is one of the
most remarkable success stories of introduced trees
to date; since its first introduction to the USA and
Conservation
Europe in 1948 it has spread to almost every coun-
This ‘conservation flagship’ conifer species oc- try with a temperate climate. Several cultivars have
curs in a limited region in the border area of two been named, especially in the Netherlands. The phe-
Chinese provinces and Chongqing Municipality notypic variation observed in planted trees obtained
(formerly E Sichuan province) in what is now inten- from early seed introductions may indicate genetic
sively cultivated and utilised countryside. It appears diversity, but trees with somewhat stunted and con-
that at least the older trees are relicts of a riparian torted lower trunks may also be due to suboptimal
Metasequoia forest, which may have stretched un- growing (climatic?) conditions. It would seem that
broken in valley bottoms before farmers moved into warm and humid summer conditions are conducive
the area only a few centuries ago (Chii & Cooper, to rapid growth, producing straight, erect trunks
1950). The several subpopulations are all reduced and long branches.
to a few to several mature trees, sometimes even a
 Microbiota Kom., Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 4: 180. 1923. Type:
Microbiota decussata Kom. (Cupressaceae).

Greek: mikros = small; bios = life; Biota is another

Description
name for Thuja; therefore a small Thuja.
Decumbent shrubs, evergreen, monoecious, spread-
ing by layering of branches, without distinct stem
Description
or trunk, up to 50–100 cm tall, often lower and
See the species description. prostrate on rocks. Bark at first orange-brown or
514 red-brown, then purplish brown, smooth, thin,
exfoliating in small flakes, turning dark brown on
Distribution
thicker stems. Branches numerous, decumbent,
As for the species. spreading and curved down in more or less plagio-
tropic frondose sprays, the whip shoots drooping,
forming a dense, spreading shrub several meters
Taxonomic notes
across. Foliage branchlets spreading at narrow angles
Recent molecular phylogenetic analysis (Gadek of ca. 20–40 degrees, slender, lax, slightly flattened,
et al., 2000) of all genera in Cupressaceae consist- ultimate branchlets to ca. 30 mm long, all covered
ently found a clade with Microbiota and Platycladus, with scale leaves. Leaves decussate, weakly dimor-
related to Tetraclinis as sister group, not to Junipe- phic, broad decurrent, imbricate, appressed but with
rus or Thuja. Phytogeographically, Microbiota and some apices free, rhombic, leaf margins mostly en-
Platycladus are nearly sympatric at present, although tire; apex acuminate or acute-pungent, 1.5–3(–3.5) ×
with a much smaller range for the former. Perhaps a 0.6–1.5 mm; abaxial stomata few, adaxial stomata in
shrubby form of the latter became isolated, due to two bands merging distally; glands central, slightly
the retreat to the south of Platycladus in Pleistocene depressed, conspicuous on facials, inconspicuous
cold phases. It could then have evolved to become on laterals; leaf colour green or yellowish green,
the decumbent shrub with minimalised cones that turning copper-brown or purplish brown in win-
is adapted to subalpine, cool and highly maritime ter. Pollen cones terminal, solitary, ovoid-globose,
mountains on the fringe of the Asian continent, 2–3 mm long; microsporophylls 10–12, decussate,
where it still occurs today in a limited area. peltate-rounded, abaxially bearing 2 relatively large,
angular-globose, yellow pollen sacs. Seed cones
somewhat hidden by foliage sprays, terminal on
Microbiota decussata Kom., Bot. Mater. Gerb. very small branchlets, developing in one season to
Glavn. Bot. Sada RSFSR 4: 180. 1923. Type: Russia: a ‘minimal’ cone of a lower small and upper larger
Russian Far East, Primoriye, Mt. Zamo-Dynza, pair of bract-scale complexes enveloping lower two
[“in monte Zamo-dinzsa”], I. K. Schischkin 158 third of single seed, spreading at maturity to 3–5 mm
(lectotype LE). Fig. 169 wide; upper pair cup-like from having surrounded
the seed, both pairs thin woody, with papillose mar-
gins and with long protruding, subapical, incurved
Etymology
bract tips. Seeds broadly ovoid, often with two lon-
The species epithet refers to the pairwise alternating gitudinal lines or ridges marked by scale edges, 2–3
opposed scale leaves. mm long; apex acutish, lustrous chestnut-brown or
blackish brown, with a lighter hilum at base.
Vernacular names
Distribution
Siberian cypress (USA); the species is also known
as Microbiota in cultivation; no common name was Russian Far East: Sikhote Alin Prov., Primoriya.
given in Flora URSS Vol. 1 (1963). TDWG codes: 31 PRM
 the Sikhote Alin Mountains, but extends northwards
Ecology
to beyond Kabarovsk. It is listed as rare in the Red
Microbiota decussata is a decumbent shrub in mon- Book of Russia and it is included in Nedoluzhko’s
tane habitats both in and outside forests, at altitudes (1999) Endangered woody plants of the Russian
between 800 m and 1200 m a.s.l. It occurs in mixed Far East. Its range is larger than the threshold for
conifer or angiosperm-conifer forest with e.g. Abies Vulnerable (VU) and its habitat is on the whole not
nephrolepis, Picea jezoensis, Pinus koraiensis, Acer threatened as it occurs in remote localities and in
ukurunduense, Alnus maximoviczii, Betula erma- vegetation types that are not subject to changes in
nii, and Sorbus amurensis. In subalpine non-forest land use, except for some forested localities nearer
vegetation it can be assosiated with Pinus pumila, Vladivostok. There is no evidence of decline.
Juniperus sabina var. davurica, or Rhododendron IUCN: LC 515
mucronulatum, often growing on talus slopes or
mossy granitic boulders with small accumulations
Uses
of soil. The climate is cold maritime with deep snow
cover for several months in winter. The ‘copper- Its decumbent habit and attractive colouring foliage
brown’ discoloring of the foliage in winter is due (green in summer and bronze in winter), combined
to chemical (anthocyanin) changes in the pigment with hardiness in cold-winter climates and easy veg-
of the mesophyll cells that may enhance the rate of etative propagation, have made this species a desir-
photosynthesis under low light conditions. able, but still not very commonly available garden
shrub. Given a well-drained, open site and cool con-
ditions, it will spread to cover the ground quickly. It
Conservation
has only been available in the trade since the 1980s
Based on available data from herbarium collections, and no cultivars are known.
this species is most common in the southern part of
 Microcachrys Hook. f., London J. Bot. 4: 149. 1845. Type: Microcachrys tetragona
(Hook.) Hook. f. [Athrotaxis tetragona Hook.] (Podocarpaceae).

Greek: mikros- = small; kachrys = catkin, cone. restricted to adaxial side of leaves, subapical, hid-
den from view. Pollen cones terminal, more or less
recurved, 2.5–4 mm long, ovoid; microsporophylls
Description
helically arranged, strongly incurved, enclosing two
See the species description. basal pollen sacs containing bisaccate or trisaccate
pollen. Seed cones usually on different branching
516 systems from pollen cones, terminal, maturing to
Distribution
6–8 mm long, ovoid, comprising of an axis with ca.
As for the species. 20 helically arranged, rounded bracts which become
fleshy, slightly swollen and red at maturity. Ovules 1
per fertile scale, inverted; seeds broadly ovoid, 1.5 ×
Microcachrys tetragona (Hook.) Hook. f., London 1.2 mm, slightly flattened when ripe, at base partly
J. Bot. 4: 149. 1845. Athrotaxis tetragona Hook., covered by the epimatium.
Icon. Pl., n.s., 2: t. 560. 1843, [“Arthrotaxis”];
Dacrydium tetragonum (Hook.) Parl., in Candolle,
Distribution
Prodr. 16 (2): 496. 1868. Type: Australia: Tasmania,
[“V. D. L.”], R. C. Gunn [367] (lectotype K). Fig. 170 Australia: Tasmania.
TDWG codes: 50 TAS
Pherosphaera hookeriana Hook. f., Fl. Tasmania 1
(5): 355. 1857, non W. Archer (1850).
Ecology
Microcachrys tetragona occurs in Tasmanian alpine
Etymology
dwarf scrubland on exposed rock outcrops, usually
The species epithet describes the four-angled above 1000 m altitude. This prostrate little shrub
branchlets due to decussate phyllotaxis of the scale spreads over rocks and through low mossy or her-
leaves. baceous vegetation in wet areas with long, whip-like
running shoots capable of rooting and ramifying to-
wards open spaces, often over bare rock. It is, despite
Vernacular names
being so low to the ground, wind pollinated as are
Creeping pine, Strawberry pine all conifers; its tiny mulberry-like red cones are suc-
culent and undoubtedly eaten by animals (birds?),
but these vectors of its seed dispersal are not known.
Description
This dwarf conifer is often growing near other,
Evergreen, monoecious or (temporarily) dioecious, taller conifers like Athrotaxis cupressoides, Diselma
prostrate shrubs 10–30 cm tall. Leading stems to ca. archeri, Pherosphaera hookeriana, Podocarpus law-
1 m long, branching frequently. Foliage branches rencei, and various angiosperms, e.g. Nothofagus
numerous, spreading or ascending; whip shoots gunnii, Epacris serpyllifolia, Orites spp., Richea spp.,
elongated, creeping out over rocks; (pen)ultimate Astelia alpina, Leptospermum ruprestre, etc.
branchlets with scale leaves quadrangular in cross-
section. Leaves on whip shoots mostly decussate,
Conservation
decurrent, lanceolate, to ca. 5 mm long, the distal
part slightly spreading, with minutely denticulate IUCN: LC
margins tapering to an acute apex. Leaves on lateral
branchlets strictly decussate, imbricate, appressed,
Uses
short triangular in appearance, 1–1.5 × 1 mm; mar-
gins minutely denticulate; dorsal side of leaves This extremely dwarfish conifer is extremely rare in
keeled towards an obtuse, incurved apex. Stomata cultivation. Despite having been introduced to the
Royal Botanic Gardens, Kew in 1862 by W. Archer, it since disappeared. In view of its natural habitat and
is still only known from a few botanical collections. climate, this species should be an interesting plant
At Kew it was grown successfully in the Temperate for the rock garden in mild winter regions where
House and in the Himalayan House, but it has long frost is uncommon.

517
 Nageia Gaertn., Fruct. Sem. Pl. 1: 191. 1788. Type: Nageia nagi (Thunb.) Kuntze
[Myrica nagi Thunb.] (Podocarpaceae).

From nagi, the vernacular name of the species that size are in some species informative, but require the
occurs in Japan. examination of sufficient samples.

1a. Leaf venation indistinct; stomata only present

Description
on the lower leaf surface 2
Dioecious or monoecious evergreen trees, rarely 1b. Leaf venation distinct; stomata present on both
518 shrubs. Numerous narrow resin canals in leaves. leaf surfaces 3
Bark thin, hard, becoming scaly. Branching irregu- 2a. Leaf blades 2–9 cm long, 0.7–3 cm wide; apex
lar, non-rhytmic, vegetative shoots terminating in acute or obtuse to truncate; pollen cones 0.5–2
small buds. Leaves spirally inserted or subopposite cm long, 2–3 mm wide; seeds always single, in-
on leading shoots, petioles twisted to position the cluding the epimatium 10–15 mm diam N. nagi
leaves in a plane, large, flat, broadly ovate-elliptic to 2b. Leaf blades 8–18 cm long, 2.5–5 cm wide; apex
lanceolate, lacking a midrib and with many paral- acute or acuminate. Pollen cones 1.5–4(–6.5)
lel, converging veins, coriaceous and more or less cm long, 3.5–4 mm wide. Seeds single or 2–3
rigid, amphistomatic or hypostomatic (stomata on together, including the epimatium 15–18 mm
abaxial side only). Pollen cones single or in small, diam N. fleuryi
spicate groups of 2–6 on axillary peduncles, ovoid- 3a. Leaf blades 2–5(–7.5) cm long, 1.5–2.5 cm wide.
cylindric, surrounded at base by small scales; micro­ Pollen cones solitary, sessile, 5–6 mm wide.
sporophylls triangular to apiculate with 2 basal, Seeds including the epimatium 13–16 mm
subglobose pollen sacs containing bisaccate pollen. diam N. motleyi
Seed cones solitary and long pedunculate, or with 3b. Leaf blades 5–30(–34) cm long, 2–9.5 cm wide.
two or more on slender shoots in the axil of a leaf Pollen cones usually in groups of 3–10, pedun-
and short pedunculate, consisting of several spirally culate, 2.5–4 mm wide. Seeds including the epi-
arranged bracts; bracts either drying up or fusing matium 15–20 mm diam 4
and enlarging at their bases, forming a weakly de- 4a. Shrubs or small trees to 10 m tall. Leaf blades
veloped, fleshy receptacle being usually only slightly (8–)16–30(–34) cm long; apex acute to acumi-
thicker than the peduncle. Only a single terminal nate. Pollen cones 2.5–3 mm wide; microsporo-
bract fertile, having an inverted ovule enveloped by phylls lanceolate. Receptacle below seed absent
the epimatium; epimatium swelling greatly, forming N. maxima
a drupe-like, fleshy to succulent, red or purple ‘fruit’ 4b. Trees to 50 m tall, rarely shrubs. Leaf blades
around the globose seed. 5–15(–23) cm long; apex acute or obtuse. Pollen
cones 3–4 mm wide, microsporophylls apicu-
5 species. late. Receptacle 12–20 mm long and succulent
red when mature N. wallichiana
Distribution
India (Assam, Kerala, Meghalaya, Nicobar Islands); Nageia fleuryi (Hickel) de Laub., Blumea 32 (1):
S China; Taiwan; Japan; Indochina (including Malay 210. 1987. Podocarpus fleuryi Hickel, Bull. Soc.
Peninsula); Malesia: from Sumatera to New Guinea Dendrol. France 76: 75. 1930; Decussocarpus fleuryi
(excluding Lesser Sunda Islands, New Britain and (Hickel) de Laub., J. Arnold Arbor. 50: 355. 1969.
New Ireland). Type not designated (syntypes J. F. Fleury 30180,
38017, E. Poilane 5963, Forest St. 8408 in P). Fig. 171
Key to the species of Nageia
Etymology
Leaf shapes and sizes are extremely variable and
these characters form a continuum from seedlings, This species commemorates the 19th century French
saplings and young trees to mature trees. Maxima in botanist J. F. Fleury.
 also in montane tropical rainforest dominated by
Vernacular names
evergreen broad-leaved angiosperm trees. It is a
kim giao nui da (Vietnamese); changye zhubai (Chi- montane species, but its altitudinal range (in Viet-
nese) nam) is somewhat broadly defined as between 500
m and 1200 m. Its large leaves indicate tolerance of
shade and it grows up well under canopy of most
Description
taller trees. As do other conifers in the main area
Trees to 25 m tall; trunk to 70 cm d.b.h. Bark scaly, of its range (northern Vietnam and adjacent south-
peeling in small, thin flakes, purplish brown; in- ern parts of China) it evades competition from an-
ner bark fibrous, reddish. Crown with ascending or giosperms by growing mostly on exposed rocky
spreading branches, pyramidal. Foliage branchlets ridges, presumably this is an edaphic adaptation us- 519
opposite (but often one of the pair not developed) ing mycorrhizal symbiosis to cope with the lack of
spreading rigidly, terete or angular, often twisted, water and nutrients. Other conifers that grow here
glabrous; terminal buds with narrowly lanceolate frequently are Pinus fenzeliana (P. kwangtungensis),
scales extending to a fine point. Leaves opposite or Pseudotsuga sinensis, and on slightly better sites
subopposite (towards end of branchlets often alter- ­Fokienia hodginsii.
nate), decussate or nearly in one plane on shaded
branches; petiole twisted ca. 90° at base, 2–5(–10)
Conservation
mm long; leaf blade variable in size and shape, 8–18
× 2.5–5 cm (largerst on young plants and shaded Widely distributed but in disjunct populations
leaves), from elliptic to ovate-lanceolate or ovate throughout NE Vietnam and SE China. Both the
with acute to acuminate apex, coriaceous, with in- extent of occurrence (EOO) and area of occupancy
distinct parallel veins, dark green above, pale green (AOO) (not calculated as herbarium specimen data
below. Stomata only on lower surface, in numerous, were considered to be incomplete) would fall out-
intermittent lines, usually conspicuous but some- side the threshold for VU. The timber is valued and
times poorly visible. Pollen cones axillary, in clusters is being selectively logged, therefore it is suspected
of 3–6, (nearly) sessile, variable in length from 1.5– that there is a global threat to this species. In Viet-
4(–6.5) cm, becoming long cylindrical at full length, nam, where this species probably has its main dis-
with a diam. of 3.5–4 mm; microsporophylls imbri- tribution, it has been assessed as Vulnerable (VU)
cate, triangular, with two pollen sacs. Seed cones according to Nguyen Tien Hiep et al. (2004). It
axillary, solitary or rarely in pairs, pedunculate, has been recorded from northern parts of Taiwan
lacking a receptacle, with a few deciduous red bracts in Flora of Taiwan (ed. 2, 2001), but these sightings
at base of fertile branchlet. Seeds single or in pairs or more likely refer to N. nagi, of which young trees
threes on fertile branches, enclosed in a light green growing in shade may have larger leaves.
(bloomed glaucous white) but when ripe purple epi- IUCN: NT
matium, the whole globose, 15–18 mm diam. when
succulent, wrinkled and dark brown when dry with
Uses
a loose seed inside.
The wood of this species has good properties, such
as straight grain, evenly texture and softness, that
Distribution
make it useful for special purposes such as fine fur-
China: Guangdong (Gaoyao, Longmen, Zengcheng), niture and musical instruments. The rather small
Guangxi (Hepu), Yunnan (Mengzi, Pingbian); Indo- stature of many trees growing in exposed situations
china: Kampuchea [Cambodia], Laos, Vietnam. makes it less valuable as a timber tree and the wood
TDWG codes: 36 CHC-YN CHS-GD CHS-GX 41 CAM is therefore only used locally. The seeds contain 36%
LAO VIE oil with a low volatility and are harvested on a mod-
est scale for this. The species has horticultural merit,
but has not been taken into cultivation widely and
Ecology
would only do well in (sub)tropical countries.
Nageia fleuryi is most often found on limestone for-
mations (karst) in mixed angiosperm/conifer ­forest;
 Nageia maxima (de Laub.) de Laub., Blumea 32 where it indeed belongs. [The genus Decussocarpus
(1): 210. 1987, [maximus”]. Decussocarpus maximus turned out to be an illegitimate name under the cur-
de Laub., J. Arnold Arbor. 50: 353. 1969. Type: rent rules of the International Code for Botanical
Malaysia: Sarawak, Sibu District, Naman F. R., Nomenclature.] De Laubenfels (op. cit.) originally
J. A. R. Anderson 3361/7 (holotype L). cited specimens from Sarawak (the type) and Su-
matera under his new species, but later [in Flora
Malesiana ser. 1, 10 (3): 394 (1988)] he omitted the
Etymology
Sumatera references, which more accurately be-
The species epithet refers to the very large leaves. long with N. wallichiana. This species remains little
known and, if more material were available, could
520 turn out to be a form with extra large leaves of
Vernacular names
N. wallichiana, whose leaves can match it in width
A very local name, ‘landin paya’, is cited in Flora if not in length.
Malesiana ser. 1, 10 (3): 394 (1988).
Distribution
Description
Borneo: Sarawak.
Shrubs or trees to 10 m tall; trunk to 20 cm d.b.h. TDWG codes: 42 BOR-SR
Bark smooth, brown; inner bark slightly fibrous,
reddish. Crown with spreading branches, bushy. Fo-
Ecology
liage branchlets opposite (but often one of the pair
not developed) spreading rigidly, terete or angular, This rare species is only known from a few localities,
often twisted, glabrous; terminal buds with narrowly one is on a ridge in moist rainforest and the oth-
lanceolate scales extending to a fine point. Leaves ers are in peat swamps. All are in lowland from near
opposite or subopposite (towards end of branchlets sea level to 120 m a.s.l. This taxon is described as a
often alternate), decussate or nearly in one plane small understorey tree and has apparently the larg-
on shaded branches; petiole twisted ca. 90° at base, est (length × width) leaves of all conifers.
4–10 mm long; leaf blade variable in size and shape,
(8–)16–30(–34) × (3–)6–9.5 cm (largerst on young
Conservation
plants and shaded leaves), from elliptic to ovate-
lanceolate with acute to acuminate apex, coriaceous, IUCN: VU (D2)
with distinct parallel veins, dark green above, pale
green below. Stomata on both surfaces, in numer-
Uses
ous, intermittent lines, usually conspicuous but
sometimes poorly visible. Pollen cones axillary, in No uses have been recorded of this species and it
clusters of 3–9, on a 3–10 mm long peduncle, cylin- does not appear to be in cultivation.
drical, 12–20 × 2.5–3 mm; microsporophylls imbri-
cate; apex lanceolate, 0.5–1 mm long, with two basal
pollen sacs. Seed cones solitary or in groups of up to Nageia motleyi (Parl.) de Laub., Blumea 32 (1):
5 on axillary branchlets, short pedunculate, lacking 210. 1987. Dammara motleyi Parl., Enum. Sem.
a receptacle, with a few deciduous bracts at base of Hort. Florent. 1862: 26. 1863; Decussocarpus motleyi
fertile branchlet. Seeds single on each bare pedun- (Parl.) de Laub., J. Arnold Arbor. 50: 352. 1969.
cle, enclosed in a green but when ripe dark purple Type: Indonesia: Borneo, Kalimantan Selatan,
epimatium, the whole globose, 15–18 mm diam. Bandjarmasin, [“Bangarmassing”], J. Motley s.n.
when succulent, wrinkled and blackish brown when (holotype K).
dry with a loose seed inside.
Etymology
Taxonomic notes
This species was named after the 19th century bota-
This species was originally described as Decusso­ nist James Motley.
carpus maximus but later transferred to Nageia,
 land “bindang-dipterocarp” forest in Sarawak, but
Vernacular names
is more often found in less tall evergreen forests on
podo kebal musang (Peninsular Malaysia); kayu podzolic sands, where it is scattered among numer-
bawa, setebal (Sumatera); medang buloh (Sarawak). ous angiosperm tree species and occasional coni-
fers. It apparently regenerates after disturbance and
is also found in secondary forest.
Description
Trees to 50 m tall; trunk to 1 m d.b.h., forming a
Conservation
straight, clear bole. Bark hard and scaly, peeling in
thin flakes, brown or reddish brown; inner bark fi- Logging of Nageia spp. in lowland forests can involve
brous, reddish. Crown with spreading branches, be- this species or the much more widely distributed 521
coming rounded. Foliage branchlets opposite (but species N. wallichiana; the ranges and ecology of
often one of the pair not developed) spreading rig- both species partly overlap in Peninsular Malaysia,
idly, terete but ultimately more or less quadrangular Sumatera and Borneo. Identification of logged trees
and often twisted, with grooves, glabrous; termi- to species is impossible if the foliage is no longer
nal buds with lanceolate scales extending to a fine available. It is possible that logging has affected this
point. Leaves opposite or subopposite (towards end species more seriously than is currently realized.
of branchlets often alternate), decussate or nearly in IUCN: LC
one plane on shaded branches; petiole twisted ca.
90° at base, 2–3 mm long; leaf blade somewhat vari-
Uses
able in size and shape, 2–5(–7.5) × 1.5–2.5 cm (larg-
erst on young plants and shaded leaves), lanceolate, Nageia motleyi is a valuable timber tree, especially
elliptic to obovate with obtuse, acute or acuminate where it grows into tall, straight trees with a long,
apex, coriaceous, with distinct parallel veins, dark clear bole. It is traded as podocarp wood. Long tim-
green above, mid green below. Stomata on both sur- ber is sawn into planks for construction (mainly
faces, in numerous, continuous or intermittent lines, house building); other uses of the wood are plywood,
conspicuous on both sides. Pollen cones axillary, veneer, interior finishing, and furniture making. As
solitary, sessile, 1.5–2 cm long, with a diam. of 5–6 it occurs within the general distribution of N. walli-
mm; microsporophylls imbricate; apex acuminate, chiana and develops into a similar tree, it is difficult
2 mm long, with two basal pollen sacs. Seed cones to assess whether uses apply to the less frequently
axillary, solitary, long pedunculate, with a few decid- encountered species or to the widespread species.
uous bracts near base of peduncle and cone bracts Nageia motleyi is probably not grown in cultivation.
fused to a swelling green then red 8–15(–20) mm
long receptacle with 5–9 exserted bract tips. Seeds at
apex of receptacle, single, enclosed in a green, when Nageia nagi (Thunb.) Kuntze, Revis. Gen. Pl. 2: 798.
ripe dark purple epimatium, the whole globose, 13– 1891. Myrica nagi Thunb., in Murray, Linn. Syst.
16 mm diam. when succulent, wrinkled and nearly Veg., ed. 14: 884. Mai-Jun 1784; Decussocarpus nagi
black when dry with a loose seed inside. (Thunb.) de Laub., J. Arnold Arbor. 50: 357. 1969.
Type not designated. Fig. 172
Distribution
Podocarpus formosensis Dummer, Gard. Chron.,
S Thailand, Borneo, Peninsular Malaysia, Sumatera.
ser. 3, 52: 295. 1912; Decussocarpus nagi (Thunb.)
TDWG codes: 41 THA 42 BOR-KA BOR-SB BOR-SR
de Laub. var. formosensis (Dummer) Silba, Phy-
MLY-PM SUM
tologia 58: 366. 1985; Nageia formosensis (Dummer)
C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: 382.
Ecology
1989; Nageia nagi (Thunb.) Kuntze var. formosensis
This species grows as tall as N. wallichiana in low- (Dummer) Silba, Phytologia 68: 38. 1990.
land tropical rainforest from ca. 15 m to ca. 500 m Podocarpus nankoensis Hayata, [Icon. Pl. Formos. 6,
a.s.l. Nageia motleyi occurs usually on dry soil but Suppl.: 74. 1917, nomen] Icon. Pl. Formos. 7: 39. 1918;
has also been found in peat swamps (“ramin-peat Nageia nankoensis (Hayata) R. R. Mill, Novon 9 (1):
swamp”) in Sarawak. It is a rare constituent of low- 77. 1999.
 Podocarpus nagi (Thunb.) Pilg. var. koshunensis when succulent, wrinkled and dark brown when dry
Kaneh., Trans. Nat. Hist. Soc. Formosa 21: 145. 1931; with a loose seed inside.
Podocarpus formosensis Dummer var. koshuensis
(Kaneh.) Merr. & Yamam., J. Soc. Trop. Agric. 7: 142.
Taxonomic notes
1935; Podocarpus koshunensis (Kaneh.) Kaneh., For-
mosan Trees, ed. 2: 36. 1936; Nageia nagi (Thunb.) Nageia formosensis (Dummer) C. N. Page was listed
Kuntze var. koshuensis (Kaneh.) D. Z. Fu, Acta Phy- in A World Checklist and Bibliography of Conifers
totax. Sin. 30 (6): 524. 1992. (Farjon, 1998, [2001]) as an accepted species. It was
by some considered a mere variety of N. nagi, while
other sources suggest that perhaps there are no
Etymology
522 fewer than three distinct species in Taiwan: Nageia
The species epithet refers to the Japanese vernacular formosensis, N. nankoensis (Hayata) R. R. Mill and
name as recorded by Carl Peter Thunberg. N. fleuryi (Hickel) de Laub. Given the variability I
have observed in specimens considered to belong
to N. nagi from mainland China, Taiwan and Japan
Vernacular names
(some of which came from planted trees), a broader
naki (Japanese); zhu bai (Chinese) concept of N. nagi is here adopted (as in Flora of
China 4, 1999), until it can be clearly demonstrated
that there are distinct and sufficient character state
Description
differences in the Taiwanese trees that merit the rec-
Trees or shrubs to 20 m tall; trunk to 50 cm d.b.h. ognition of one or more separate species, or even
Bark scaly, peeling in small, thin flakes, reddish or varieties of N. nagi.
purplish brown, weathering grey. Crown with as-
cending or spreading branches, becoming rounded,
Distribution
or bushy. Foliage branchlets opposite (but often one
of the pair not developed) spreading rigidly, more S China: Fujian, Guangdong, Guangxi, Hainan,
or less tetragonal, often twisted, with grooves in Hunan, Jiangxi, Sichuan, Zhejiang; Japan: Kyushu,
the edges, glabrous; terminal buds with narrowly Nansei-Shoto (Ryukyu Is.), Shikoku; Taiwan.
lanceolate scales extending to a fine point. Leaves TDWG codes: 36 CHS 38 JAP-KY JAP-SH NNS TAI
opposite or subopposite (towards end of branch-
lets often alternate), decussate or nearly in one
Ecology
plane on shaded branches; petiole twisted ca. 90°
at base, 3–8(–15) mm long; leaf blade variable in Nageia nagi occurs in mixed mesophytic evergreen
size and shape, 2–9 × 0.7–3 cm (largerst on young forest and mixed mesophytic deciduous forest
plants and shaded leaves), from narrowly elliptic (Wang, 1961). It occurs in hills and low mountains
to ovate-lanceolate or obovate with acute to obtuse from about 200 m to 1200 m a.s.l. In evergreen oak
or nearly truncate apex, coriaceous, with indistinct forest it is one of several shade tolerant conifers that
parallel veins, dark green above, pale green below. may occur under canopy or take advantage of small
Stomata only on lower surface, in numerous, inter- gaps to break through: Taxus chinensis, Cephalo-
mittent lines, usually conspicuous but sometimes taxus fortunei, Keteleeria fortunei, and Fokienia
poorly visible. Pollen cones axillary, in clusters of hodginsii are the most common of these. Besides
up to 10 or sometimes solitary, (nearly) sessile, vari- Castanopsis spp. and Quercus spp. (the oaks), nu-
able in length from 0.5–2 cm, becoming cylindrical merous angiosperm trees occur in these forests or
at full length, with a diam. of 2–3 mm; microsporo­ forest remnants. In Taiwan and southern Japan the
phylls imbricate, apiculate, with two pollen sacs. coniferous element of this vegetation is more domi-
Seed cones axillary, solitary and long pedunculate, nant, with Pseudotsuga sinensis or P. japonica and
or rarely in pairs, with a few deciduous bracts near Tsuga sieboldii often added to the mixture. In forest
base of fertile branchlet and some remaining dried or woodland on drier mountain slopes N. nagi tends
bracts subtending a single seed enclosed in a green, to follow streams, but it is known to regenerate in
when ripe dark purple epimatium, usually bloomed more open thickets after forest disturbance.
white, the whole (sub)globose and 10–15 mm diam.
 Conservation Vernacular names
This species is widely distributed, but it is almost Due to its wide range numerous local names have
impossible to establish from herbarium collection been recorded; Flora Malesiana 1, 10 (3): 393 (1988)
data where it is truly indigenous (growing in the lists many for the Malesian region; in China it is
wild) and where it has been introduced and planted. known as rou tuo zhu bai and in Vietnam it is called
Wilson (1916) noted that he did not meet with any kim giao nui dat.
trees growing in the wild whilst traveling in Japan.
In the wild, it is limited to a few forest remnants
Description
in the most southern parts of that country, includ-
ing the oceanic Ryukyu Islands. One can take the Trees (rarely shrubs) to 50 m tall; trunk to 1m d.b.h., 523
view (as the Chinese and Japanese mostly do), that forming a straight, clear bole. Bark hard and scaly,
such a taxon is not threatened with extinction even peeling in thin flakes, dark brown or reddish brown;
if its wild (sub)populations are greatly reduced. In inner bark 5–6 mm thick, slightly fibrous, pinkish or
Taiwan, S. Y. Lu (1996) has assessed the species as reddish. Crown with spreading branches, becoming
Critically Endangered (CR), with wild growing trees rounded. Foliage branchlets opposite (but often one
restricted to Taipei Co. in the north and Taitung Co. of the pair not developed) spreading rigidly, terete
in the far south of the island. Similar inventories but ultimately slightly flattened and often twisted,
based on intimate knowledge of the forest flora are with grooves, glabrous; terminal buds with lanceo-
lacking for most of mainland China. late scales extending to a fine point. Leaves opposite
IUCN: DD or subopposite (towards end of branchlets often al-
ternate), decussate or nearly in one plane on shaded
branches; petiole twisted ca. 90° at base, 5–10(–15)
Uses
mm long; leaf blade extremely variable in size and
Nageia nagi is a valuable timber tree, but its most shape, 5–15(–23) × 2–6(–9) cm (largerst on young
common use is as an amenity tree in China and Ja- plants and shaded leaves), from narrowly elliptic
pan, where it is found in many of the climatically to ovate-lanceolate, ovate or falcate (some terminal
milder parts of these countries planted in gardens, leaves) with acute to obtuse apex, coriaceous, with
parks, sanctuaries, and even as street trees. It is also distinct parallel veins, dark green above, mid green
popular as a tree for bonsai cultivation. It is much below. Stomata on both surfaces, in numerous, in-
less commomly planted in Europe, the USA and termittent lines, usually conspicuous on underside
New Zealand, where it is almost restricted to botani- and less so on upperside. Pollen cones axillary, in
cal collections. clusters of up to 7–10 on a 4–10 mm long peduncle,
variable in length from 0.8–2 cm, becoming cylin-
drical at full length, with a diam. of 3–4 mm; micro­
Nageia wallichiana (Presl) Kuntze, Revis. Gen. sporophylls imbricate, apiculate, with two pollen
Pl. 2: 800. 1891. Podocarpus wallichianus C. Presl, sacs. Seed cones axillary, solitary, long pedunculate,
Abh. Königl. Böhm. Ges. Wiss., ser. 5, 3: 540. 1846; with a few deciduous bracts near base of peduncle
Decussocarpus wallichianus (C. Presl) de Laub., and cone bracts fused to a swelling green then red
J. Arnold Arbor. 50: 349. 1969; Podocarpus latifolius to purplish red 12–20 mm long receptacle with 4–7
Wall., Pl. Asiat. Rar. 1 (2): 26, t. 30. 1830, non Mirb. exserted bract tips. Seeds at apex of receptacle, sin-
(1825); Nageia latifolia (Wall.) Gordon, Pinetum: gle, enclosed in a green, when ripe dark purple epi-
138. 1858. Type: India: Pundu Mts., Mt. Sillet, [“in matium, the whole globose, 15–20 mm diam. when
montibus Punduae”], N. Wallich 6050 (holotype succulent, wrinkled and nearly black when dry, with
K-W). Fig. 173, 174 a loose seed inside.

Etymology Distribution
This species was named after Nathaniel Wallich China: Yunnan; India: Assam, Kerala (Nilgiri and
(1786–1854), who was an early student of the Indian Palani Hills), Andaman Islands, Nicobar Islands;
flora.
 I­ ndochina; Malesia (but not in Central & E Jawa and ergreen subtropical forest dominated by Castanopsis
on the Lesser Sunda Islands only on Flores). and/or Quercus on hillsides but not in high moun-
TDWG codes: 36 CHC-YN 40 ASS-AS ASS-ME IND- tains. In margins of peat swamps, in mossy forest
KE 41 AND-AN CBD LAO MYA NCB THA VIE 42 on sandstone plateaus of Sarawak, and on mountain
BOR-BR BOR-KA BOR-SB BOR-SR JAW LSI-LS MLY- ridges with clay or sand amongst rocks it becomes
PM MOL PHI SUL SUM stunted. In New Guinea it is sometimes associated
with Araucaria and Podo­carpus in mixed conifer
forests, which also have several species of Fagaceae,
Ecology
especially in the genus Castanopsis.
This is the most widespread species in the genus
524 Nageia and perhaps also one of the most truly tropi-
Conservation
cal of all conifers, as it occurs near sea level in dip-
terocarp forest on the equator. It is scattered but IUCN: LC
often common in primary rainforest with canopy
heights to 50 m or more, and occurs from lowlands
Uses
to montane forested ridges at 2100 m a.s.l. In the
lowland rainforest it develops a straight bole lifting Nageia wallichiana is a highly valued timber tree, es-
its crown into the canopy. It is, however, not a long- pecially where it grows into tall, straight trees with
lived emergent and boles usually are rather slender a long, clear bole. It is traded as podocarp wood.
without buttresses indicating modest longevity. Un- Long timber is sawn into planks for construction
like Agathis (Araucariaceae) it is not gregarious. (mainly house building); other uses of the wood are
Kerangas (forest on leached sandy soils) can have plywood, veneer, interior finishing, furniture mak-
both species, as well as other conifers like Dacry- ing, and sometimes (Fly River, Wagu, Papua New
dium, Dacrycarpus, Falcatifolium falciforme, and Guinea) the construction of small canoes. Small
Sundacarpus amarus, mixed with Myrtaceae and stems are used for household utensils, drumsticks,
other angiosperms that have adapted to poor nutri- etc. It is not grown in cultivation other than in a few
ent situations. In China, N. wallichiana occurs in ev- botanic gardens.
Neocallitropsis Florin, Palaeontographica B 85 (1–8): 590. 1944. Type: Neocallitropsis
pancheri (Carrière) de Laub. [Eutacta pancheri Carrière] (Cupressaceae).

Callitropsis R. H. Compton, J. Linn. Soc., Bot. 45: a conical or more often a rounded or flat-topped
432. 1922, non Oerst. (1864). Type: Callitropsis arau- open ‘candelabra’ crown. Foliage branchlets in
carioides R. H. Compton [Neocallitropsis pancheri dense tufts of 20–30, crowded at the terminal 10–15
(Carrière) de Laub.] cm of main branches, assurgent or erect, sparsely
branched or unbranched, to 20 cm long, 5–10(–12)
Latin: neo- = new; Callitropsis (genus name) means mm wide, densely covered with imbricate leaves,
‘similar to Callitris’; the prefix is here used to avoid mostly determinate and deciduous, some continu- 525
a later homonym. ous. Leaves in alternate whorls of 4, whorls turned
45 degrees around shoot axis relative to the previ-
ous whorl, seemingly in 8 rows, short decurrent or
Description
nearly sessile with broad base, lanceolate, incurved,
See the species description. thick, coriaceous, rigid, keeled, 6–15 × 1.8–2.5 mm;
margins minutely serrate; apex acute-pungent; sto-
mata in two bands abaxially, in more scattered rows
Distribution
adaxially; leaf colour yellowish green or green. Pol-
As for the species. len cones terminal, 10–12 × 6–7 mm, subglobose to
ovoid; microsporophylls in 3–4 alternating whorls of
4, 3–6 × 2 mm, peltate-rhombic with a long, acute
Neocallitropsis pancheri (Carrière) de Laub., Fl. or acuminate apex, bearing (2–)6–14 abaxial, small
Nouv. Calédonie Dépend. 4: 161. 1972. Eutacta pollen sacs. Seed cones terminal on short branchlets,
pancheri Carrière, Traité Gén. Conif., ed. 2, 2: 864 maturing within a year to an opened cone up to 15
[615]. 1867. Type: New Caledonia: Grande Terre, mm long with spreading bract-scale complexes. Ma-
Province Sud, Montagnes de Yaté, E. Vieillard 1274 ture bract-scales in two alternating whorls of 4, with
(holotype P). Fig. 175, 176 lower scales slightly larger than upper ones, 10–12 ×
3 mm and 8–10 × 2 mm, subulate to linear, similar
Callitropsis araucarioides R. H. Compton, J. Linn. to mature foliage leaves but with adaxial thicken-
Soc., Bot. 45: 432. 1922; Neocallitropsis araucarioides ing, leaving margins of outer bracts and apices un-
(R. H. Compton) Florin, Palaeontographica B 85 altered but reflexed, rostrate; margins of thickened
(1–8): 590. 1944. tissue papillose. Columella short pyramidal, 1.5 mm
tall. Seeds 1–2(–4) per cone, angular-ovoid, acutish,
6–7 × 2.5–3.5 mm, light brown, with 2(–3) marginal
Etymology
0.5–0.7 mm wide wings.
The species epithet commemorates the French bota-
nist Jean A. I. Pancher (1814–1877).
Distribution
New Caledonia: Province Sud (SE part), one loca-
Vernacular names
tion in Province Nord (Mt. Paéoua).
No common names have been recorded for this spe- TDWG codes: 60 NWC
cies.
Ecology
Description
In scrubland (‘maquis minier’) on ultramafic rock
Shrubs or small evergreen trees to 4–6(–10) m tall, (serpentine, ‘cuirasse maquis’), following stream
multistemmed or monopodial, stem to 30–50 cm courses and along lower mountain ridges up to about
diam. Bark on large stems exfoliating in narrow 950 m a.s.l., rarely higher. This species is often associ-
strips, grey. Branches spreading wide or ascending, ated along streams with the podocarps Dacrydium
higher order branches assurgent or erect, forming araucarioides, D. guillauminii, and ­Dacrycarpus
 vieillardii, elsewhere with Agathis ovata, Calli- now virtually ceased. No assessment of any decline
tris neocaledonica, Podocarpus novae-caledoniae, due to this exploitation has come to the attention of
and angiosperms, including many shrubs as well the Conifer Specialist Group of IUCN-SSC, yet the
as ­Cyperaceae. Several of the trees and shrubs also slow growth of this species makes it likely that this
develop the candelabra-like crowns typical for low has occurred. A large population covering ca. 122 ha
trees and shrubs in this environment, but grow more is protected in the Montagne des Sources (Parq de
solitary than Neocallitropsis, which is strongly gre- la Rivière Bleue) and smaller populations are under
garious. The climate is tropical with abundant rain- protection at the Chûte de la Madeleine (17.5 ha) and
fall through most of the year. in the Plaine des Lacs (3.5 ha). Despite this protec-
tion, continued decline and fragmentation of popu-
526 lations are projected.
Conservation
IUCN: EN [A2c, B1ab(iii)+2ab(iii)]
The habitat of this species is extremely susceptible
to fire and wildfires are a widespread hazard in New
Uses
Caledonia. With less than 10 populations known in
the southern mountains and one, recently discov- The resin in the wood of this species is used to make
ered, in the northern part of Grande Terre, con- an oil extract, which is sold as ‘Araucaria oil’; this is
servation issues are apparent. In the past, there was now only locally traded. It is also locally used as an
unsustainable exploitation of the wood of this slow ornamental shrub, but does not appear to have been
growing conifer for oil extraction, but this use has taken into cultivation outside New Caledonia.
A HANDB O OK OF T H E WOR L D’ S C ON I F E R S
Giant Sequoia (Sequoia gigantea) Drawing by Aljos Farjon
A HANDB O OK OF THE WORLD’ S C ONIFE RS

by

AL JO S FA R JON

Volum e I I

BRILL
Leiden-BOSTON
2010
 This book is printed on acid-free paper.
Library of Congress Cataloging-in-Publication Data
The Library of Congress Cataloging-in-Publication Data is available from the Publisher.

front cover:
Cunninghamia konishii foliage

back cover:
top left: Pinus wallichiana seed cones
bottom right: Cephalotaxus haringtonii ripe seeds

isbn: 978 90 04 17718 5

© Copyright 2010 by Koninklijke Brill NV, Leiden, The Netherlands.

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Nothotsuga Hu ex C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: 390. 1989. Type:
Nothotsuga longibracteata (W. C. Cheng) Hu ex C. N. Page [Tsuga longibracteata
W. C. Cheng] (Pinaceae).

Greek: nothos = bastard, base-born; i.e. a (putative) otherwise remote, with oblique, twisted petiole,
hybrid between Tsuga and an unnamed genus. 1.1–2.4 cm long, 1–2(–2.5) mm wide, linear, abruptly
tapering at both ends; apex acutish, faintly grooved
above, slightly flattened; margins entire; stomata
Description
5–10 lines along the median groove above, two
See the species description. bands separated by a faint midrib below (abaxial 533
side); leaf colour dark glossy green, stomatal bands
whitish green. Pollen cones subterminal on small
Distribution
lateral shoots, clustered in umbels from a single
As for the species. bud, pedunculate, pendant, 5–10 mm long, yellow-
ish brown (in sicco). Seed cones lateral, or subtermi-
nal on small shoots, more or less erect on 5–10 mm
Nothotsuga longibracteata (W. C. Cheng) Hu ex long peduncles, ovoid oblong to cylindrical, with
C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: obtuse truncate apex, (2–)2.5–5(–5.8) cm long, 1.5–
390. 1989. Tsuga longibracteata W. C. Cheng, Contr. 2.5(–3) cm wide with opened scales, purplish or red
Biol. Lab. Sci. Soc. China, Sect. Bot. 7 (1): 1. 1932. when immature, ripening to dark brown; old cones
Type: China: Guizhou, Yinjiang Tujiazu Miaozu persisting several years, breaking off at peduncle,
Zizhix, Fanjing Shan, [“Yinkiang”], Y. Tsiang 7712 or sometimes disintegrating; cone rachis blackish
(holotype NAS). Pl. 22, Fig. 177, 178 brown. Seed scales 20–30, suborbicular to broadly
peltate-auriculate, convex, opening slightly or very
wide (reflexed), 1–2.2 × 1.2–2.5 cm at mid-cone;
Etymology
abaxial surface sparsely pubescent when immature,
The species epithet describes the relatively long soon glabrous, finely striated; upper margin round-
bracts on the seed cone. ed, entire or erose; base short pedicellate. Bracts
subspathulate, erose-denticulate at acute apex, 7–18
mm long, straight, not exserted beyond margins of
Vernacular names
more distal seed scales. Seeds ovoid, 4 × 2.5–3 mm,
Bristlecone Hemlock; changbao tieshan (Chinese) brown; seed wings ovate-oblong, 7–12 × 5–6 mm,
reddish brown.
Description
Taxonomic notes
Trees to 30 m tall, d.b.h. to 1–1.2 m; trunk monopo-
dial, often forked or multi stemmed; bark soon flak- This taxon was first described as a species of Tsuga
ing, rough and scaly, brownish grey. Branches long, by the well-known Chinese botanist W. C. Cheng in
heavy, curved, ascending or spreading; crown of 1932. Hu (1951) proposed a separate genus Notho­
young trees conical, with a drooping leader, of old tsuga for this species, but failed to give a Latin de-
trees open and irregular or dense and flat topped. scription; the genus name was then validated by
Branchlets slender, firm, not drooping, brown- Page (1989). French botanists in the ‘School of Gaus-
ish yellow to brown, in 2nd or 3rd year grey; mostly sen’ at Toulouse proposed a generic hybrid origin
glabrous, rarely minutely pubescent, with weakly between Keteleeria and Tsuga, but gave no evidence
developed pulvini; on 2–3 year old twigs develop for this and applied an illegitimate name. Chinese
numerous small, lateral, leaved shoots (5–15 mm), botanists (e.g. Flora of China 4: 39–40, 1999) do not
which do not extend much further. Vegetative buds recognize its status as a distinct genus, but there are
ovoid, acutish at apex, 2–4 mm long, not resinous, several distinctive characters in both male and fe-
shining brown. Leaves more or less pectinate, on male cones not shared by other species of Tsuga in
small lateral shoots in false whorls on emergence, Asia or North America that appear to justify generic
534

7
4

10

5
2
7 8
11

Pl ate 2 2 . Nothotsuga longibracteata. 1. Habit of trees. 2. Branch with foliage. 3, 4. Seed cones. 5. Seed cone
scales. 6. Bract. 7, 8. Leaves. 9. Seeds. 10. Short shoot with leaves. 11. Pollen cones.
recognition. Its phylogenetic position based on both Notho­tsuga longibracteata and Tsuga chinensis. Pinus
morphological and DNA data confirms this taxon- massoniana or P. fenzeliana (syn. P. kwangtungensis)
omy. Despite its name, there is no evidence that this locally dominate the general canopy of broad-leaved
taxon is of hybrid origin. trees on poorer sites, where N. longibracteata is also
concentrated.
Distribution
Conservation
China: S Fujian, N Guangdong, NE Guangxi, NE
Guizhou, SW Hunan, Jiangxi, NW Yunnan. This species is considered to be Endangered, be-
TDWG codes: 36 CHC-GZ CHC-YN CHS-FJ CHS-GD cause it is very rare despite its relatively wide distri-
CHS-GX CHS-HN CHS-JX bution. Large scale logging has depleted the number 535
of trees to an unquantified extent (Fu & Jin, 1992)
and substantial parts of forest with this species have
Ecology
gone, especially at lower elevations. In 1982 N. longi­
Nothotsuga longibracteata occurs on low to medi- bracteata was found near Gongshan in the far NW
um high mountains, at elevations between 300 and of Yunnan at 3200 m a.s.l. by a Chinese team; I veri-
2300(–3200) m a.s.l. It grows on both red and yellow fied its correct identity in the Beijing Herbarium
earth. The climate is humid and warm-temperate (PE) in October 2000. This disjunct occurrence at
to wet and cool, with annual precipitation between high altitude could indicate that its actual distribu-
1000–2000 mm. The species occurs in two forest for- tion remains incompletely known.
mations (Wang, 1961). In the evergreen broad-leaved IUCN: EN (A1c)
forest formation mostly with sclero­phyllous broad-
leaved trees such as Castanopsis spp., Lithocarpus
Uses
spp., Quercus spp., and with ­Fokienia hodginsii; in
the deciduous mixed mesophytic forest at higher In China this species is considered to be a desirable
elevations with Fagus longipetiolata, Tetracentron forest tree suitable for afforestation. Its use as a tim-
sinensis, Nyssa sinensis, Acer angustilobium, Davidia ber tree must be limited due to its rarity. It is not
involucrata, Sorbus spp., etc. In the evergreen broad- in general cultivation outside China and rare in bo-
leaved forest formation there are stands of pure tanical collections.
 Papuacedrus H. L. Li, J. Arnold Arbor. 34: 25. 1953. Type: Papuacedrus papuana
(F. Muell.) H. L. Li [Libocedrus papuana F. Muell.] (Cupressaceae).

Papua = New Guinea; Cedrus is the classical name c­ arinate, cuspidate; laterals bilaterally flattened, 2–3
for (true) cedars. mm long on old, ‘mature’ foliage, up to 20 mm long
on whip shoots of young plants, lanceolate to ob-
long; margins entire, with incurved, appressed or
Description
free, obtuse or acute apex; few stomata on upperside
See the species description. and 2 broad primary stomatal bands on underside;
536 gland absent or inconspicuous at base of facials; leaf
colour lustrous olive green to dark green, with glau-
Distribution
cous green stomatal bands. Pollen cones terminal,
As for the species. solitary, cylindrical, 6–25 × 2–3 mm; microsporo-
phylls 8–30, decussate or in whorls of 4, peltate, with
acute or rounded apex, bearing (2–)4(–6) abaxial
Papuacedrus papuana (F. Muell.) H. L. Li, J. Arnold pollen sacs near lower margin. Seed cones (sub)
Arbor. 34: 25. 1953. terminal, on 2–10 mm long branchlets with rhom-
bic, acute scale leaves, thin woody, 8–18 × 5–8 mm
when closed, changing from green to glaucous green
Etymology
to brown or dark blackish brown. Bract-scale com-
The species epithet refers to Papua, a collective name plexes consisting of 2 decussate pairs; lower pair 4–7
for the island of New Guinea and the Australasian × 2–5 mm, curved, widest at base; upper pair much
people who were its first settlers. larger, 7–17 × 3–7 mm, elliptic, widest in the middle
section where the recurved, acute bract tip emerges,
distal part slightly reflexed, truncate or obtuse, ru-
Vernacular names
gose with grooves radiating from bract tip abaxially,
De Laubenfels, in Flora Malesiana 1, 10 (3): 446 smooth or striate adaxially, with whitish seed scars
(1988), cites numerous local names applied in New near base. Seeds 2–4, angular-ovoid or oblique, 2–5
Guinea to this species. × 1–3 mm, red-brown with a whitish hilum; wings
2 on opposite sides, thin membranous; largest wing
4–7 × 2–5 mm; smallest wing often reduced to a strip
Description
1–2 mm wide, translucent yellowish brown.
Trees to 20(–50) m tall, evergreen, monoecious, of-
ten appearing dioecious; trunk normally monopo-
Taxonomic notes
dial, slender, to 60 cm d.b.h. Bark often spirally
twisted, exfoliating in shaggy scales or strips; outer The morphological differences in the leaves between
bark light brown, weathering grey. Branches short Papuacedrus papuana var. papuana and P. papuana
and spreading in sheltered (younger) trees, long, var. arfakensis are minor and disappear with the
ascending to nearly erect in high montane forest, ‘maturation’ of the foliage. As in many other conifers,
forming conical to pyramidal crowns, or ± umbel- leaves in a juvenile stage differ markedly in shape and
late, flat-topped crowns at higher altitudes. Foliage size from leaves in an adult stage and, as in this case,
branches mostly in flattened sprays, denser and as- there are often transitional forms as well. The ranges
surgent in older and exposed crowns, branching of both varieties overlap, but var. arfakensis only
distichous, frondose, ultimate branchlets gradu- occurs in the western parts of New Guinea and is
ally shorter, entirely covered with flattened leaves. presumably the only variety in the Moluccas, where
Leaves on lateral branchlets scale-like, in whorls the species remains undercollected (Johns, 1995).
of 4 or decussate, strongly dimorphic, imbricate; From herbarium specimens with adult-type foliage
facials much smaller than laterals, 1 mm on old, and without pollen cones it becomes impossible to
‘mature’ foliage, up to 8 mm long on whip shoots of determine whether the sample represents ­either one
young plants, rhombic to lanceolate in appearance, of the varieties. Van Royen (1979), perhaps with this
difficulty in mind, recognized one species without or dance grounds of villages (Johns, 1995). In horti-
any infraspecific distinctions. culture it is only known from a few botanic gardens.

2 varieties are recognized:

Distribution
Malesia: Maluku [Moluccas]; Papuasia: New Guin- Papuacedrus papuana (F. Muell.) H. L. Li var.
ea. papuana. Libocedrus papuana F. Muell., Trans.
TDWG codes: 42 MOL 43 NWG-IJ NWG-PN Roy. Soc. Victoria 1 (2): 32. 1889. Type: Papua New
Guinea: Owen Stanley Range, Mt. Knutsford,
[probably Owalama Range], W. MacGregor 286
Ecology
(holotype MEL). Fig. 179, 180 537
This species, as it occurs along a substantial altitudi-
nal gradient, is present in different forest zones from
Description
montane tropical rainforest to subalpine scrubland.
The altitudinal range is (620–)900–3600(–3800) m Lateral pair of transitional leaves on young plants
a.s.l.; its greatest extent and abundance is reached in spreading widely from base up to 6 mm at the far-
the mossy cloud forest zone from ca. 1500 m to the thest point below apex, falcately curved; apex turned
tree line. In the lower montane rainforest it is a scat- upwards or slightly recurved; intermediate leaves
tered emergent associated with angiosperms such similar but less widely spreading. Pollen cones 6–15
as Casuarina, Castanopsis, Cinnamomum, Engel­ mm long, with up to 16 decussate or whorled micro-
hardtia, Halfordia, Lithocarpus, Schizomeria, and sporophylls.
Xanthostemon; in higher montane forest it becomes
more prevalent with Nothofagus, Cryptocarya, and
Distribution
Eugenia. It can also form conifer dominated forest,
occasionally with Araucaria cunninghamii var. pa­ New Guinea.
puana, but more often with Dacrydium spp., Phyllo­ TDWG codes: 43 NWG-IJ NWG-PN
cladus hypophyllus, and Podocarpus spp. These small
conifer forests are often surrounded by fire-induced
Conservation
grasslands (Imperata cylindrica) with a mantle zone
of ericaceous shrubs, or by tree fern grassland. Many Despite logging activities in large parts of its range
of these conifers are often present in lower densities that have been associated with human habitation for
in the lower montane rain forests as well. At lower a very long time, this variety is still abundant and
altitudes Papuacedrus grows mostly on basic soils. not considered threatened with extinction.
In mossy forest, along high mountain streams and IUCN: LC
in subalpine scrubland the trees are stunted and the
soils are often acidic and water-logged; at around Papuacedrus papuana (F. Muell.) H. L. Li var.
3000 m or higher swamps are often surounded arfakensis (Gibbs) R. J. Johns, Curtis’s Bot. Mag.
by Papuacedrus-Phyllocladus-Podocarpus woodland 12 (2): 70. 1995. Libocedrus arfakensis Gibbs,
with tussocks of Gahnia dominant in the under­ Contr. Phytogeogr. Fl. Arfak Mts.: 84, f. 6a-b. 1917;
storey (Johns, 1995). Rainfall is generally high but Papuacedrus arfakensis (Gibbs) H. L. Li, J. Arnold
seasonal, with up to 4000 mm per year on the high- Arbor. 34: 25. 1953; Libocedrus papuana F. Muell.
est slopes. var. arfakensis (Gibbs) de Laub., Fl. Malesiana, ser.
1, 10 (3): 446. 1988. Type: Indonesia: Papua, Arfak
Mts., Koebre Ridge, L. S. Gibbs 5594 (lectotype BM).
Uses
The timber of this species is widely used for con-
Description
struction (mainly building of houses in villages); in
some areas the fibrous bark is used for roof covering Lateral pair of transitional leaves on young plants
and insulating house walls. It is rarely cultivated, al- spreading slightly from base up to 3 mm at the far-
though individual trees may be planted at ‘sing-sing’ thest point below apex, apically recurved towards
base of the following pair; intermediate leaves like
 adult leaves but larger. Pollen cones 15–25 mm long, 160,929 km2, falls well outside the threshold for a
with up to 30 whorled microsporophylls. threatened category. However, it is also disjunct,
with two localities in Maluku (Moluccas) and one
around the Wissel Lakes in New Guinea. The tim-
Distribution
ber of this tree is widely used for the construction of
Malesia: Maluku [Moluccas] (Bacan and Obi Is- houses in rural villages. It is therefore suspected that
lands); Papuasia: New Guinea: Papua (Vogelkop some decline may have occurred in more densely
Pen­insula, Sudirman Mts.). populated areas such as around the Wissel Lakes. Its
TDWG codes: 42 MOL 43 NWG-IJ most widespread stands are probably in the Arfak
Mountains. It is appropriate to flag this variety as
538 near threatened.
Conservation
IUCN: NT
The known range of this variety is relatively wide
and its extent of occurrence (EOO), calculated at
Parasitaxus de Laub., Fl. Nouv. Calédonie Dépend. 4: 44. 1972. Type: Parasitaxus
usta (Vieill.) de Laub. [Dacrydium ustum Vieill.] (Podocarpaceae).

Latin: parasitus = a parasite; Taxus is the classical mm long, 1–2 mm wide (on older branches larger
Latin name for yew. [While now understood as a and being forced apart by the thickening branch),
distinct family, podocarps were formerly classified broadly lanceolate to more or less triangular; apex
with Taxaceae]. appressed or free, obtuse or acute. Stomata on both
sides of leaves, conspicuous and scattered on abax-
ial side. Pollen cones usually on the same branch-
Description
ing systems as seed cones, terminal, solitary, more 539
See the species description. or less oval, 3–3.5 mm long, 1.5–2 mm wide; micro-
sporophylls 8–13, imbricate, broadly triangular with
erose upper margin, terminating in an incurved
Distribution
apex, with two basal pollen sacs. Fertile ovulate
As for the species. branchlets numerous, erect. Seed cones mostly ter-
minal, some lateral, consisting of 3–6 spreading nar-
row and apically incurved red bracts of which the
Parasitaxus usta (Vieill.) de Laub., Fl. Nouv. Calé- distal 1–2 subtend short pedunculate fertile scales
donie Dépend. 4: 45. 1972, [“ustus”]. Dacrydium bearing a single terminal and inverted ovule. Young
ustum Vieill., Ann. Sci. Nat. Bot., sér. 4, 16: 56. 1861; seeds ovoid-­oblong with an apical crest, when ma-
Podocarpus ustus (Vieill.) Brongn. & Gris, Bull. ture completely surrounded by a globose, hard,
Soc. Bot. France 13: 426. 1866; Nageia usta (Vieill.) glaucous white epimatium, the whole 3–4 mm diam.
Kuntze, Revis. Gen. Pl. 2: 800. 1891. Type: New Cal-
edonia: Grande Terre, Province Sud, Poila, (moun-
Taxonomic notes
tains around Poila), E. Vieillard 1267 (holotype P).
Fig. 181, 182 This remarkable little conifer was first described
as a species of Dacrydium in 1861 and a few years
later transferred to the genus Podocarpus and then,
Etymology
in 1891, incomprehensibly at least under the present
Usta is a Latin word for a red pigment and here re- circumscription of that genus, to Nageia. Not until
fers to the red colour of the branchlets. [The ending 1959, when David de Laubenfels published a brief
of the epithet changes with the gender of the genus note on it in the journal Science, was it demonstrat-
name: -a = feminine, -um = neuter, -us = mascu- ed to be parasitic on another conifer in the Podo­
line.] carpaceae, Falcatifolium taxoides. It is the first and
only parasitic gymnosperm ever recorded. Its spe-
cies epithet under Dacrydium (neuter) was ustum,
Vernacular names
under Podocarpus (masculine) ustus, but under
Cèdre rabougri (NC French) ­Parasitaxus, which may not be a Taxus taxonomi-
cally but certainly is so for purposes of Latin no-
menclature, it has to be usta (feminine).
Description
Small erect monoecious shrubs to 150 cm tall but
Distribution
usually less than 100 cm, multi-stemmed or some-
times with a single stem. Bark thin, with large lenti- New Caledonia (Grande Terre).
cels, on lower stem breaking into small grey scales. TDWG codes: 60 NWC
Branches numerous, spreading and ascending, con-
torted, the higher order branchlets covered in red-
Ecology
dish scale leaves, the (pen)ultimate branchlets 2–3
mm thick, scaly with leaves, purple. Leaves scale- Parasitaxus usta is the only known parasitic gymno-
like, spirally arranged, imbricate and decurrent, 2–3 sperm. Its host is another podocarpaceous ­conifer,
 Falcatifolium taxoides, a small tree. The parasite at- in the Montagnes des Sources and another in the
taches to the roots (sometimes the [underground] mountain range culminating in Mont Panié in the
stem base) of the host, but how the seed of P. usta north. Its actual abundance is difficult to ascertain
germinates and how the seedling makes the connec- due to its cryptic habit in dense, shady undergrowth;
tion is still not fully understood. The most likely sce- most individuals are being observed close to estab-
nario involves the help of a fungus (Deutero­mycotina lished mountain trails. Its host is a small tree not ex-
or ‘fungi imperfecti’), which establishes a mycelian ploited for timber. Due to its shunning of sunlight,
endophyte that enters the root cambium of the host closed-canopy primary forest is essential habitat for
and the haustorial base of the parasite (Woltz et this strange conifer, and local deforestation may al-
al., 1995). Plants of P. usta are nearly always found ready have reduced populations and would threaten
540 growing (seemingly in the litter-covered soil) under others. It is present in a number of well protected
trees of F. taxoides, but sometimes on the base of the mountain forest areas, including the Montagnes des
trunk. The leaves of P. usta contain chlorophyll and Sources and Mont Panié.
are stomatiferous, but are not green and its capacity IUCN: VU [B1ab(iii)+2ab(iii); C2a(i)]
of independent carbon fixation is severely reduced.
Its habitat is therefore that of its host: moist tropical
Uses
montane angiosperm forest with scattered conifers,
commonly with emergent Araucaria spp. or less of- There are no uses recorded of this unique species
ten Agathis spp. Parasitaxus usta grows usually in and attempts to cultivate it in some botanic gardens
the deep shade of a multi-layered canopy, never in have thus far remained unsuccessful. It is a sacred
full sunlight. plant in the tradition of the Kanaks, the first people
to colonize New Caledonia from Melanesia.
Conservation
Parasitaxus usta is found scattered across the main
island (Grande Terre), with a concentration of sites
Pherosphaera W. Archer, Hooker’s J. Bot. Kew Gard. Misc. 2: 52. 1850, p.p., quoad
descr. foem. Type: Pherosphaera hookeriana W. Archer (Podocarpaceae).

Microstrobos J. Garden & L. A. S. Johnson, Contr. Pherosphaera fitzgeraldii (F. Muell.) Hook. f.,
New South Wales Natl. Herb. 1 (6): 315. 1951. Type: Hooker’s Icon. Pl. 4: t. 1383. 1882. Dacrydium fitz­
Microstrobos fitzgeraldii (F. Muell.) J. Garden & geraldii F. Muell., Fragm. 11: 102. 1880; Microstrobos
L. A. S. Johnson fitzgeraldii (F. Muell.) J. Garden & L. A. S. Johnson,
Contr. New South Wales Natl. Herb. 1 (6): 316. 1951.
Greek: phero = to bear; sphaira = ball, globe; refer- Type: Australia, New South Wales, Blue Mountains,
ring to the shape of the seed cones. R. Fitzgerald s.n., 1880, 1881 (syntypes, ?MEL, n.v.). 541

Description Etymology
Dioecious evergreen shrubs. Resin cavities in leaves. This species was named by Ferdinand von Mueller
Bark smooth or rough and scaly. Branches spreading after R. Fitzgerald, who presented him with the first
and rigid or pendulous and creeping. Adult and ju- botanical specimens.
venile leaves similar, spirally arranged, 2–4(–6) mm
long and mostly scale-like. Stomata on adaxial side, Vernacular names
hidden from view. Pollen cones terminal, globose
Dwarf mountain pine
to ovoid, 2–6 mm long; microsporophylls (6–)8–15,
spirally arranged, with two basal pollen sacs; pol-
Description
len with 2–3 air sacs. Seed cones terminal on small
branchlets, more or less globular, 2–4 mm long, Ascending, erect or drooping shrub to 1 m tall and
with (3–)5–8 spreading, fertile scales, not fleshy at 2 m wide. Bark smooth, brown, weathering grey.
maturity. Mature seeds usually 1–4 per cone, solitary Branches slender, drooping to pendulous or strag-
and basal on adaxial side of a fertile scale (bract), gling over rocks; foliage branches very lax and slen-
exposed, erect, ovoid with a slightly constricted and der, irregularly branched two to four times. Leaves
truncated apex. monomorphic, spirally arranged, decurrent at
base, the free part spreading, 2–4 mm long (vary-
2 species. ing in length on a single branchlet), ca. 1 mm wide
(wider at base, narrower at apex), subulate, apically
incurved, keeled on abaxial side, convex on adax-
Distribution
ial side; apex obtuse or apiculate; leaf colour dark
Australia: New South Wales, Tasmania. olive green, whitish on adaxial side where stomata
are situated. Pollen cones terminal, solitary, globose
maturing to ovoid, 4–6 × 3 mm; microsporophylls
Key to the species of Pherosphaera
10–15, spirally arranged, ovate-oblong with erose-
– Branches drooping to pendulous; foliage denticulate margin and two basal pollen sacs. Seed
branches very lax and slender; leaves 2–4 × 1 cones terminal and solitary on erect branchlets,
mm P. fitzgeraldii more or less globular, 2–4 mm long, with 4–8 fertile,
– Branches spreading, contorted; foliage branch- spreading, broadly lanceolate to ovate bracts. Ma-
es assurgent or spreading; leaves 1 × 1 mm ture seeds usually 1–3 per cone, solitary and basal on
P. hookeriana adaxial side of a fertile scale (bract), exposed, erect,
ovoid with a slightly constricted and truncated apex,
ca. 1 × 0.7 mm, dull brown.

Distribution
Australia: New South Wales (Blue Mountains, from
Katoomba Falls to Wentworth Falls).
 TDWG codes: 50 NSW-NS Pherosphaera hookeriana W. Archer, Hooker’s
J. Bot. Kew Gard. Misc. 2: 52. 1850, p.p., excl. typ.
Dacrydium hookerianum (W. Archer) Eichler,
Ecology
in Engler & Prantl, Nat. Pflanzenfam. 2 (1): 107.
Pherosphaera fitzgeraldii has an extremely limited 1887. Type: Australia: Tasmania, [“V. D. Land”],
distribution and specific habitat. This species occurs R. C. Gunn s.n. (lectotype K). Fig. 183
on the sides (on ledges) and at the bottom of water-
falls which cascade off the sandstone escarpment on Microstrobos niphophilus J. Garden & L. A. S. John-
the eastern side of the Blue Mountains. These falls son, Contr. New South Wales Natl. Herb. 1 (6): 316.
have fluctuating amounts of water dependent on 1951, non rite publ. (Art. 36); Pherosphaera nipho­
542 precipitation levels that have occurred on the pla- phila (J. Garden & L. A. S. Johnson) Florin, Taxon 5
teaus above. The altitude where the species grows (8): 191. 1956, non rite publ. (Art. 36).
ranges from 600 to 930 m a.s.l. This shrubby, often
trailing species is only found within the spray zone
Etymology
of the waterfalls. It occurs there with other shrubs,
ferns and mosses, in part under tree canopy, in part This species was named after the English botanist
in the open but then often shaded part of the day as Joseph Dalton Hooker (1817–1911), the second Di-
it predominantly grows on S-facing precipices. In- rector at the Royal Botanic Gardens, Kew.
vasive weeds (Hedera, Rubus) occur among several
populations.
Vernacular names
Mount Mawson pine
Conservation
This species is extremely rare and has suffered a sub-
Description
stantial decline. Baker & Smith (1910) wrote: “at the
base of most of the chief falls on the Blue Moun- Erect, densely branched shrub to 2.5 m tall. Bark
tains” but it is now known to grow only on a few on larger stems rough, exfoliating with small scales,
cliffs in the Wentworth Falls and Katoomba Falls, an brown weathering blackish grey. Branches spread-
extent of occurrence ca. 9 km long. A survey in 1988 ing, contorted; foliage branches assurgent or spread-
reported 455 individual plants in 7 populations. Five ing, divided up to four times until very small; (pen)
of these populations were on government land (Blue ultimate branchlets 1–1.5 mm thick, forming dense
Mountains National Park), two on private land. and stiff tufts of foliage. Leaves spirally arranged,
Recruitment is very low or almost negligible and a scale-like, in seedlings and young plants short lan-
slow decline has been observed directly over several ceolate, 2–2.5 × 1 mm, largely free but incurved,
years. Threats are or have been urban development, keeled, replaced higher up the plant by smaller
water pollution, habitat degradation, and competi- leaves; in mature plants all imbricate and closely ap-
tion from invasive species (Hedera helix, Rubus sp.), pressed, broadly triangular, mostly ca. 1 × 1 mm, on
as well as native shrubs and trees. whip shoots to 2.5 × 1.5 mm, on older, thicker branch-
IUCN: EN (C2a (i)) lets to 2 × 2 mm, concave with a blunt keel; margins
minutely serrate; apex obtuse; leaf colour yellowish
green. Stomata on adaxial side, hidden from view.
Uses
Pollen cones numerous, terminal, globose, ca. 2 mm
Dwarf mountain pine is rare in cultivation, but diam. colour red-brown at anthesis; microsporo-
makes an attractive rock-dwelling small shrub. It is phylls (6–)8–12, spirally arranged, rounded with
presently mostly confined to botanic gardens, but serrulate margins and two basal pollen sacs. Seed
should be suitable for rock gardens close to water cones terminal on distally down-curved branchlets,
features including fountains. Its growth in cultiva- more or less globular, 3–4 mm long, with (3–)5–8
tion would also contribute substantially to ex situ spreading, fertile, ovate and concave scales with an
conservation of this endangered species. acute apex, browning at maturity. Mature seeds usu-
ally 1–4 per cone, solitary and basal on adaxial side
of a fertile scale (bract), exposed, erect, ovoid with
a slightly constricted and truncated apex, ca. 1.3 × often associated with Athrotaxis cupressoides, Micro­
1 mm, lustrous brown. cachrys tetragona and, usually on somewhat drier
sites, with Diselma archeri; frequent angiosperms
are Nothofagus gunneri, Richea pandanifolia, R. sco­
Distribution
paria, and Eucalyptus coccifera, while cushion form-
Australia: Tasmania (central and western moun- ing peat mosses (Sphagnum) cover the ground in
tains). many places.
TDWG codes: 50 TAS
Conservation
Ecology
IUCN: LC 543
Pherosphaera hookeriana occurs in the subalpine
regions of the Tasmanian highlands, usually above
Uses
1000 m a.s.l. It is frequent in wet moors and often
fringes the lakes and tarns that are numerous in This shrubby species is rare in cultivation, being
these mountains. Precipitation is high and occurs grown in some specialist nurseries and often con-
year-round; temperatures are cool with sleet and fused with Diselma archeri (Cupressaceae). It should
snow falling in most months of the year, but mostly be hardy in areas with mild and wet winters where
in winter. There is no extended period of snow cover frost is light. It will make a suitable shrub for rock
as the climate is extremely oceanic. Bedrocks are gardens or for pot-grown balcony or patio ever-
acidic granites, gabbro, and gneiss and the waters greens.
have a low pH of 4.5–5 on average. This species is
 Phyllocladus Rich. ex Mirb., Mém. Mus. Hist. Nat. 13: 48. 1825 (nom. cons.). Type:
Phyllocladus aspleniifolius (Labill.) Hook. f. [Phyllocladus billardieri Mirb. (nom.
illeg.) (Podocarpus aspleniifolius Labill.)] (Phyllocladaceae).

Brownetera Rich. ex Tratt., Ann. Mus. Natl. Hist. Distribution

Nat. (Paris) 16: 299. 1810. Type: Brownetera asple­
Malesia: Borneo, Sulawesi, Philippines, Maluku
niifolia (Labill.) Tratt. [Podocarpus aspleniifolius
[Moluccas], New Guinea; Australia: Tasmania; SW
Labill.] Thalamia Spreng., Anleit. Kennt. Gewächse,
Pacific: New Zealand.
ed. 2, 2: 218. 1817. Type: Thalamia aspleniifolia (La-
544 bill.) Spreng. [Podocarpus aspleniifolius Labill.]
Key to the species of Phyllocladus
Greek: phyllos = leaf; klados = branch, shoot; in bo- Reproductive organs (pollen and seed cones) are not
tanical Latin: phyllocladium = a leaf-like branch or always present but have some diagnostic characters
shoot. necessary for correct determination. As a further
aide therefore the country or region of origin is also
given with each species.
Description
Shrubs or trees, usually dioecious, occasionally 1a. Phylloclades simple or pinnately compound;
monoecious, evergreen. Resin in bark and phyllo- margins crenately or obtusely lobed. 2
clades. Bark smooth, becoming scaly. Branches in 1b. Phylloclades pinnately compound; margins en-
pseudo-whorls at intervals along a main or single tire to deeply dissected 3
stem, plagiotropic and/or ascending (Rauh’s model); 2a. Phylloclades mostly simple. Pollen cones on a
lateral branches of highest order transformed to short branching system, solitary or with 2–5 to-
simple or compound (pinnate) phylloclades (green gether. Native of Tasmania P. aspleniifolius
planated branchlets of variable size and shape) ar- 2b. Phylloclades pinnately compound or simple.
ranged spirally or in pseudo-whorls. Indetermi- Pollen cones from a terminal bud with up to 10
nate shoots ending in a globose bud with imbricate in a pseudo-whorl. Native of New Zealand
scales. True leaves on seedlings on the main stem, P. trichomanoides
linear, changing gradually to subulate and scaly, in- 3a. Seed cones 4–8(–10) alternately on either side
conspicuous and deciduous on lateral branchlets in of the petiolate phylloclade base. Pollen cones
older plants. Stomata in mature plants restricted to from a terminal bud with 15–20 in a pseudo-
phylloclades. Pollen cones crowded in a low spiral at whorl. Native of New Zealand P. toatoa
or below apex of leading normal shoots, becoming 3b. Seed cones on the margins or in an apical notch
more or less remotely placed with shoot elongation, of phylloclades, solitary or with 2–3 together.
subtended by perular scales, stalked, cylindrical; Pollen cones on a short branching system, up
microsporophylls with a small triangular head and to 15 together. Native throughout Malesia
two relatively large pollen sacs containing bisaccate P. hypophyllus
pollen. Seed cones (sub)marginal or terminal on
petiolate or foliate parts of phylloclades, in rows, in Phyllocladus aspleniifolius (Labill.) Hook. f.,
pairs or solitary; consisting of a few to many bract London J. Bot. 4: 151. 1845. Podocarpus aspleniifolius
scales, of which 1-several are fertile; ovules axillary Labill., Nov. Holl. Pl. Sp. 2: 71, t. 221. 1806. Type:
to a bract scale, solitary, erect, subtended by an aril, Australia: Tasmania, North Esk River, Cataract
which arises after the ovule has been formed. Seeds Gorge, [“Habitat in capite Van-Diemen”], J. J. H. de
ovoid, dorsiventrally compressed, partly embedded Labillardière s.n. (lectotype FI). Fig. 184
by the fused, swollen and succulent bract scales of
the cone and subtended by a filmy white aril leaving
Etymology
the apical part free. Seedlings with 2 cotyledons.
The species epithet compares this species with the
4 species. foliage of Asplenium ruta-muraria, a small Europe-
an fern common on rocks and ancient walls.
 of several bracts, 2–5 of which are fertile and merge
Vernacular names
to a red or purplish structure 3–5 mm long, which
Celery-top pine becomes slightly swollen and pinkish red (drying
leathery brown). Seeds 2–5 per cone, 1 per fertile
bract, each in a white aril covering the lower two
Description
third of seed; distal part free, 5 mm long, semi-ovoid
Shrubs or trees to 20 m tall, in tall forest with a (laterally flattened) with a lateral ridge and a small
clear bole to 10 m or more, d.b.h. to 50 cm. Bark to protrusion at apex, greenish black to black.
20 mm thick, with large lenticels; outer bark deeply
furrowed and scaly in old trees, exfoliating in small
Distribution
to medium sized flakes, dark brown weathering 545
dark grey or blackish; inner bark close to wood Australia: Tasmania.
red or pink, slightly fibrous. Branches spreading TDWG codes: 50 TAS
or ascending, forming a narrow or wide pyramidal
crown. Foliage branches mostly straight, spreading
Ecology
at an angle of less than 90°, robust , terete, smooth,
newest shoots tinged red, becoming green then light Phyllocladus aspleniifolius occurs in montane tem-
brown, terminating in a short bud with spreading, perate rainforest up to the tree line from near sea
triangular to acicular scales. True leaves on seed- level on the west coast of Tasmania to 1200 m a.s.l.
lings 10–15 mm long, subulate-linear, 1 mm wide, in the central highlands. The largest trees grow in
with a midvein and stomata on abaxial (lower) side, mixed forest with Eucalyptus spp. at lower eleva-
acute; along new shoots and on margins of phyllo­ tions. At higher altitude (above 700–800 m a.s.l.) it
clades filiform leaves appear in young plants, 1–3 is found in open woodland with Eucalyptus cocci­
mm long, deciduous. Phylloclades axillary to re- fera, Nothofagus cunninghamii, N. gunnii, Richea
duced, deciduous, filiform scale leaves, mostly sim- pandani­folia, R. scoparia, Athrotaxis cupressoides,
ple, bifacially flattened, leaf-like, (1.5–)2.5–5(–8) cm A. selaginoides, A. ×laxifolia, and various shrubs.
long, mostly cuneate or rhombic in outline, with Towards the tree line it grows with Orites acicula­
crenately to obtusely lobed margins, narrowing to ris, O. revoluta, Tasmannia lanceolata, Podocarpus
a cuneate or petiolate base, sometimes more or less lawrencei, Diselma archeri, Pherosphaera hookeri­
pinnatifid; phylloclades on old trees smallest and ana, Nothofagus gunnii, Olearia spp., and heath-like
least dissected, on seedlings often pinnatisect. Vena- dwarf shrubs and alpine herbs. The substrate is
tion of phyllo­clades penni-parallel, with a midvein acidic and derived from dolerite, granite, or quar-
from base to (near) apex and few to numerous veins zite, and is usually well drained, such as thin soil on
running parallel to each other from midvein or mid- boulder or scree slopes. Precipitation is abundant
vein line to margins under a narrow but sometimes through much of the year, with no marked periods
widening angle (i.e. they curve outwards). Flushing of drought.
phylloclades reddish or rusty brown; new phyllo-
clades bright green or tinged red; old phylloclades
Conservation
lustrous deep green or dark green above, pale green
below, sometimes glaucous. Stomata numerous on IUCN: LC
the underside (abaxial) in irregular lines. Pollen
cones on a short branching system from a terminal
Uses
bud, solitary or with up to 5 cones together, with one
or two small bracts (‘foliola’) at their base, cylindri- Celery-top pine varies from a medium size tree in
cal, 5–8 mm long, 2–2.5 mm wide, pink or reddish the forests at middle altitudes to a shrub in the sub-
when immature, becoming yellow. Microsporo- alpine zone. The wood of good size trees is straight
phylls ovoid-triangular, with two basal, globose pol- grained and dense, pale brown, and not dissimilar
len sacs. Seed cones axillary to reduced scale leaves to yew (Taxus). It is used for construction, flooring,
on margins or at base of reduced phylloclades, or on ship masts, furniture, and cabinet work. This species
terminal short branching systems without phyllo­ is rare in cultivation, limited to botanic gardens and
clades, solitary or with 2–4 together, each consisting arboreta and a few private gardens.
 Phyllocladus hypophyllus Hook. f., Icon. Pl., n.s., 5: under a ­narrow but sometimes widening angle (i.e.
t. 889. 1852. Type: Malaysia: Sabah, Ranau District, they curve outwards). Flushing phylloclades yellow-
Mt. Kinabalu N. P., H. Low s.n. (holotype K). Fig. green, red, or rusty brown; new phylloclades bright
185, 186 green or tinged red or copper, sometimes glaucous;
old phylloclades lustrous deep green or dark green
above, pale green below, sometimes glaucous. Sto-
Etymology
mata numerous on the underside (abaxial) in ir-
The species epithet is composed of Greek hypo = be- regular lines. Pollen cones at base of a new shoot or
low or under and phyllus = leaved. from a terminal large bud, usually on a branching
system with up to 15 cones, each cone on a 5–25 mm
546 long peduncle subtended by a strap-like, scarious
Vernacular names
bract, sometimes with a few reduced phylloclades
Celery pine, Celery-top pine; bindang, pelayo (Bor- and with two small bracts (‘foliola’) at their base,
neo-Sarawak); kayu karongan, rapak-rapak (Bor- cylindrical, 10–15 mm long, 3 mm wide, pink or
neo-Kalimantan); kayu empire (Sulawesi); bejalin reddish when immature, becoming yellow. Micro-
(Maluku); dalung (Pilippines). sporophylls ovoid-triangular, often acuminate, with
two basal, globose pollen sacs. Seed cones axillary to
reduced scale leaves on margins of phylloclades or
Description
in an apical notch, or on terminal short branching
Shrubs or trees to 40 m tall, in tall forest with a clear systems without phylloclades or with a few much
bole to 20 m or more, d.b.h. to 100 cm, usually more reduced phylloclades, solitary or with 2–3 together,
slender. Bark to 25 mm thick, with large lenticels; each consisting of several bracts, 1–3 of which are
outer bark becoming scaly, exfoliating in small to fertile and merge to a red or purplish cupulate struc-
medium sized flakes, dark brown weathering dark ture 5–7 mm long, which becomes slightly swollen
grey or blackish; inner bark close to wood red or and bright red (drying leathery brown). Seeds soli-
pink, slightly fibrous. Branches spreading or ascend- tary to each fertile bract, with a white or yellow aril
ing, forming a narrow or wide crown much depend- covering the lower half of seed only, 5–7 mm long,
ing on location. Foliage branches mostly straight, semi-ovoid (laterally flattened) with a small protru-
spreading at an angle of less than 90°, robust, terete, sion at apex, ripening to lustrous tan or chestnut
smooth, newest shoots tinged red, becoming green brown.
then light brown, terminating in a short bud with
spreading, triangular to acicular scales. True leaves
Distribution
on seedlings 5–10 mm long, subulate-linear, 0.5–0.7
mm wide, with a midvein and stomata on abax- Malesia: Borneo, Maluku [Moluccas], Philippines,
ial (lower) side, acute; at base of phylloclades and Sulawesi; Papuasia: New Guinea.
on their margins filiform leaves appear in young TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
plants, 2–5 mm long, deciduous. Phylloclades ax- MOL PHI SUL 43 NWG-IJ NWG-PN
illary to reduced, deciduous, filiform scale leaves,
pinnately compound, the pinnate segments bifa-
Ecology
cially flattened, leaf-like, extremely variable in size
and shape, 1–13 cm long, 0.5–5 cm wide, generally Phyllocladus hypophyllus occurs in lower montane
rhombic in outline but also parabolic to lanceolate, to subalpine evergreen rainforests at altitudes be-
with entire to deeply dissected margins (crenate tween (310–)600 m and 3400(–4000) m a.s.l. At
to laciniate), the latter type mostly on seedlings to lower altitudes it grows as a canopy tree of consid-
saplings, narrowing to a petiolate base; phylloclades erable size with other conifers, e.g. Agathis sp. in
with seed cones smaller than sterile ones, irregularly kerangas on white sand derived from sandstone, or
dissected, with few lobes, often cuneiform or deeply in mixed forests with Podocarpaceae, Fagaceae and
emarginate. Venation of phylloclade segments pen- Lauraceae as the dominant families of trees. It is
ni-parallel, with a midvein from base to (near) apex also found in high montane cloud forest or ‘mossy’
and few to numerous veins running parallel to each forest, which remains lower than 20 m and is char-
other from the midvein or midvein line to margins acterized by epiphytic growth of ferns and mosses.
Conifers, including P. hypophyllus, Dacrydium sp.,
Description
Dacrycarpus sp., and Podocarpus sp. may dominate,
or these forests are mixed with angiosperms. In New Small trees to 15 m tall, d.b.h. to 60 cm. Bark with
Guinea Nothofagus grandis is often the dominant large lenticels; outer bark becoming fissured and
tree species, with Phyllocladus and podocarps mixed scaly, exfoliating in small to medium sized flakes,
in. At still higher altitudes the forest is dwarfed and dark brown weathering dark grey or blackish.
P. hypophyllus becomes shrubby, often growing on Branches ascending, forming a narrow or wide py-
the edges of boggy grasslands (especially in New ramidal or rounded crown. Foliage branches mostly
Guinea) or on rocky ridges. This species is found on straight, spreading at an angle of less than 90°, ro-
a variety of substrates, such as granite, serpentine, bust, terete, smooth, newest shoots green then light
sandstone, peaty soils, and sometimes volcanic de- brown, terminating in a short bud with free, trian- 547
posits or eroded limestone. gular, acuminate and dorsally keeled scales. True
leaves on seedlings 5–10 mm long, linear, 0.5–0.7
Conservation mm wide, acute, often interspersed with juvenile,
deeply pinnatifid, 3–5 cm long phylloclades; at base
IUCN: LC
of phylloclades and on their margins filiform leaves
appear in young plants, 2–5 mm long, deciduous.
Uses
Phylloclades axillary to reduced, deciduous, filiform
As a timber tree this species is of local importance scale leaves, pinnately compound, pinnate segments
only due to its scarcity as a big forest tree. In Pa- bifacially flattened, leaf-like, 1.5–6 cm long, 1–4 cm
pua New Guinea, a ban on the export of its unsawn wide, rhombic to obovate-flabellate in outline, with
timber is in force in order to stimulate domestic entire to deeply dissected margins (crenate to lac-
processing. The wood is very similar to that of Podo­ iniate on seedlings), narrowing to a subsessile to
carpus and straight grained, fine textured and easy petiolate base. Venation of phylloclade segments
to work, but non-durable for outdoor purposes. It is penni-parallel, with a midvein from base to half
used for light construction, flooring, interior finish, length and few to numerous veins running nearly
joinery, cupboards, and to a limited extent for the parallel to each other to margins (i.e they gradu-
making of furniture. It has excellent properties for ally curve outwards). Flushing phylloclades copper
plywood and veneer. More specialized uses are for coloured; new phylloclades lustrous bright green
laboratory bench tops, foundry patterns and stor- or glaucous green; old phylloclades with a bronze
age batteries. The resin (copal) has been collected tinge. Stomata numerous on underside (abaxial) in
by tapping the trees. The bark is used for roofing in irregular lines. Pollen cones at base of a new shoot
New Guinea and the phylloclades are used to make or from a terminal large bud, in pseudo-whorls of
tea in Borneo. This species is in cultivation, but rare up to 15–20 cones, each cone on a 10–25 mm long,
and mainly limited to botanic gardens. stout peduncle subtended by a strap-like, scari-
ous bract, cylindrical, 10–20(–25) mm long, 3–4
mm wide, with two small bracts (‘foliola’) at base,
Phyllocladus toatoa Molloy, New Zealand J. Bot. greenish to lemon when immature, becoming yel-
34: 290. 1996. Phyllocladus glaucus Kirk, Trans. low. Microsporophylls ovoid-triangular, with two
New Zealand Inst. 1: 149. 1869 [‘glauca’], non basal, globose pollen sacs. Seed cones 4–8(–10) al-
Carrière (1855). Type: New Zealand: North Island, ternately on either side of the long petiolate base of
Great Barrier Island, Kiwiriki, T. Kirk WELT 37785 fertile phylloclades, on short stalks and subtended
(holotype WELT). by a single ligulate bract, consisting of 10–20 spirally
arranged, imbricate bracts, which swell slightly at
maturity and turn from green with purple tips to
Etymology
lustrous purplish green. Seeds 8–20 per cone, 1 to
Toatoa is the Maori name for this species. each fertile bract, with a white aril mostly hidden
below bracts and covering the lower half of seed
only, 3–4 mm long, semi-ovoid (laterally flattened)
Vernacular names
with a faint lateral ridge, purple ripening to blackish
Celery-top pine; toatoa (Maori) brown, sometimes tan.
 climate regions, but should be an ideal small tree for
Taxonomic notes
horticulture with its bright bluish young phyllodes
This species is still known in many books on New of new growth.
Zealand trees as Phyllocladus glaucus, or wrongly
spelled ‘glauca’, but it has been demonstrated by
Brian Molloy (1996) that this name was wrongly ap- Phyllocladus trichomanoides D. Don, in Lambert,
plied by Kirk as a later homonym of Carrière’s name Descr. Pinus, ed. 8°, 2: 159. 1832.
(1855) to a different taxon. Carrière described a live
plant in cultivation at the Jardin des Plantes in Paris,
Etymology
which in all likelyhood was just a glaucous form of
548 P. aspleniifolius from Tasmania. The New Zealand The species epithet (Latin trichoma = hair) means
taxon was still unknown to European botanists in ‘hair-like’ and refers to the shape of true leaves in
1855 and in the second edition of Carrière’s book Phyllocladus.
(1867) the author states that P. glaucus [‘glauca’] was
introduced from Tasmania to France in 1853. Conse-
Vernacular names
quently, the taxon known by its Maori name toatoa
was in need of a new species name, for which Molloy Celery pine; tanekaha (Maori); Mountain toatoa
chose to use the native name. (Maori in part)

Distribution Description
New Zealand: North Island (on or N of the 39° S Shrubs or trees to 23 m tall, in tall forest with a clear
parallel). bole to 10 m or more, d.b.h. to 100 cm, at high alti-
TDWG codes: 51 NZN tudes a spreading shrub. Bark to 25 mm thick, with
large lenticels; outer bark smooth, becoming scaly
in old trees, exfoliating in small flakes, dark brown
Ecology
weathering grey with blackish patches; inner bark
Phyllocladus toatoa occurs in mixed subtropical to close to wood red or pink, slightly fibrous. Branches
warm temperate rainforest from near sea level to spreading or ascending, forming a narrow or wide
600 m a.s.l. It is a minor constituent of the kauri pyramidal crown. Foliage branches mostly straight,
forest (Agathis australis) which is dominated by spreading at an angle of less than 70°, robust , terete,
conifers, with as many as eight species per hectare smooth, green turning light brown, terminating in
in some places. In podocarp-hardwood forest where a short bud with spreading, triangular scales. True
angiosperms usually dominate, it will be restricted leaves on seedlings 10–15 mm long, filiform to subu-
to poorer sites. Much of this forest has gone and the late-linear, 1 mm wide, with a midvein and stomata
remnants are often opened up by past logging of big- on abaxial (lower) side, acute; along new shoots and
ger trees, leaving a more open secondary growth in margins of phylloclades subulate leaves may appear,
which P. toatoa, being a sub-canopy tree in mature 1–4 mm long, semi-deciduous. Phylloclades axil-
tall forest, has a better chance to thrive. lary to reduced, deciduous, subulate scale leaves,
pinnately compound or simple, bifacially flat-
tened, leaf-like, (0.5–)2–8(–12) cm long, (segments)
Conservation
cuneate, rhombic or trullate in outline, with crenate
IUCN: LC to obtusely lobed margins, narrowing to a cuneate
or petiolate base; phylloclades on old trees smallest
and least dissected, on seedlings pinnatifid or pin-
Uses
nately compound. Venation of phylloclade segments
The wood of toatoa or celery-top pine is nearly penni-parallel, with a midvein (or axis) from base
white, straight grained and strong; it is used for fur- to (near) apex and few to numerous veins running
niture, but being rare and now protected from com- parallel to each other from midvein or midvein line
mercial exploitation not of economic importance. It to margins under a narrow but sometimes widening
is only cultivated in some botanic gardens in mild angle (i.e. they curve outwards). New phylloclades
bright green or tinged red; old phylloclades lustrous
Distribution
deep green or dark green above, pale green below,
sometimes glaucous. Stomata numerous on the New Zealand: North Island, South Island.
underside (abaxial) in irregular lines. Pollen cones TDWG codes: 51 NZN NZS
from a terminal bud, in semi-whorls of up to 10
cones together, sessile or on 3–10 mm long pedun-
Ecology
cles, with one or two small bracts (‘foliola’) at their
base, cylindrical, 8–10 mm long, 2.5–3 mm wide, Phyllocladus trichomanoides occurs in lowland to
purple to brick red when immature, becoming red- subalpine forests and woodlands, from near sea
brown with yellow pollen sacs. Microsporophylls level to 1800 m a.s.l. In mixed lowland conifer for-
ovoid-triangular, acute, with two basal, globose pol- ests Phyllocladus trichomanoides is growing with 549
len sacs. Seed cones axillary to reduced and decidu- Agathis australis in Northland and with Phyllocladus
ous scale leaves on the margin or apex of reduced toatoa, Prumnopitys ferruginea, Podocarpus totara,
phylloclade segments, or at base of undivided phyl- and Dacrydium cupressinum. In areas with highest
loclades, solitary or with 2–4 together, each consist- rainfall there is usually a mixed angiosperm-conifer
ing of several bracts, 2–3 of which are fertile and forest with e.g. Metrosideros umbellata, Weinmannia
merge to a reddish or purplish green structure 3–5 racemosa, and Quintinia acutifolia as the dominant
mm long, which becomes slightly swollen and pur- angiosperms and, besides the already mentioned
ple (drying brown). Seeds 2–3 per cone, 1 per fertile conifers, P. trichomanoides can here be associated
bract, each in a white aril covering lower two third with Dacrycarpus dacrydioides, Prumnopitys taxi­
or more of seed; distal part free, 5 mm long, semi- folia and Manoao colensoi.
ovoid (laterally flattened) with a lateral ridge and a
small protrusion at apex, greenish black to black.
Uses
The wood of Celery pine is straight grained, heavy,
Taxonomic notes
and strong and used for construction, carpentry,
Keng (1978) keyed out the species in Phyllocladus ship masts, mine props, furniture making, and cabi-
based on the distinction between simple and pin- net work. Some of this wood is nicely figured with
nate phylloclades, with the result that he classified reddish, yellowish, and whitish and resembles yew
Phyllocladus alpinus Hook. f. as a variety of the (Taxus); it is excellent for parket floors and cabi-
Tasmanian species P. aspleniifolius. This taxonomic nets. Native forests are now mostly protected from
treatment has been followed by subsequent authors. exploitation, so the avaialble quantities of Celery
However, it would imply a rather unlikely occur- pine wood are now very limited. The Maoris made
rence of one species on two disparate island groups a red dye with the use of tannins contained in the
that have been widely separated for millions of years. bark and this can be used with certain types of soft
New Zealand and Tasmania share some genera of leather. In cultivation Celery pine is not uncommon
conifers, both extant and extinct (Farjon, 2008: 173) in some countries with a mild winter climate, espe-
but no species. The distinction between the two cially Cornwall and Ireland, where it can be seen in
types of phylloclades is actually less sharp and, as large gardens and parks. The shrubby alpine form is
Keng himself acknowledged, forms a morphologi- more hardy and is also in cultivation.
cal continuum, with both forms occurring in all
species (Tomlinson & Takaso, 1989). In similar high 2 varieties are recognized:
montane to subalpine habitats, the phylloclades (or
their segments) are likely to have converged in their Phyllocladus trichomanoides D. Don var. tricho­
shapes between the New Zealand and Tasmanian manoides. Type not designated.
species. The reproductive organs of P. alpinus are
similar to those of P. trichomanoides, but differ in
Description
more characters from those of P. aspleniifolius. It is
for these reasons that P. alpinus is here classified as Trees to 23 m tall. Phylloclades mostly pinnately
a variety of P. trichomanoides, which leaves two spe- compound or pinnatifid. Pollen cones with 5–10
cies in New Zealand and one species in Tasmania. mm long peduncles.
 Phyllocladus trichomanoides D. Don var. alpinus
Distribution
(Hook. f.) Parl., in Candolle, Prodr. 16 (2): 498.
New Zealand: North Island, N South Island. 1868. Phyllocladus alpinus Hook. f., Fl. Nov.-Zel. 1:
TDWG codes: 51 NZN NZS 235, t. 53. 1853; Phyllocladus aspleniifolius (Labill.)
Hook. f. var. alpinus (Hook. f.) H. Keng, J. Arnold
Arbor. 59: 263. 1978. Type: New Zealand: South
Ecology
Island, Nelson, J. C. Bidwill 137; [locality unknown],
Var. trichomanoides is found as far south as the lectotype K, here designated). Fig. 187
northern end of South Island and is from there re-
placed by the more ubiquitous var. alpinus. At higher
Description
550 altitudes var. trichomanoides grows with Libo­cedrus
bidwillii, Podocarpus cunninghamii, Halocarpus bi­ Shrubs or small trees to 9 m tall with a conical or py-
formis, and various angiosperm shrubs and small ramidal habit. Phylloclades mostly simple, shallowly
trees. It can form dense thickets interspersed with or deeply lobed. Pollen cones on short peduncles.
tussock grassland and dwarf shrubs. It is also a
component of subalpine beech forest (Nothofagus
Distribution
solandri), which can vary from pure beech to mixed
beech-conifer forest. New Zealand: North Island, South Island.
TDWG codes: 51 NZN NZS
Conservation
Ecology
IUCN: LC
Var. alpinus is found from ca. 500 m a.s.l. upwards,
but in the far south and west of South Island it is
found at sea level as well.

Conservation
IUCN: LC
Picea A. Dietr., Fl. Gegend Berlin 2: 794. 1824. Type: Picea abies (L.) H. Karst. [Pinus
abies L.] (Pinaceae).

Veitchia Lindl., Gard. Chron. 1861: 265. 1861. (nom.

Synopsis
rej.). Type: Veitchia japonica Lindl. [incertae sedis].
The classifications of the genus Picea since those of
Picea is the classical name for spruces. Willkomm (1887) and Mayr (1890) up to Liu (1982),
have exclusively or primarily the vegetative characters
of the species as their basis and virtually ignore the im-
Description
portant characters of the female cones. They can only 551
Monoecious evergreen trees with a monopodi- be interpreted as ‘artificial’. More recently, Schmidt
al, straight trunk. Resin canals in the wood, bark, (1989) has given a classification primarily based on
leaves and seed cones. Branching in rhythmic pseu- cone characters, with the characters of the leaves as
do-whorls at regular intervals on trunk, largely pla- a secondary criterion. Schmidt’s system is largely fol-
giotropic, sometimes pendulous (Massart’s model). lowed here, but with the rank of subgenus equivalent to
Bark usually flaking with papery scales, becoming the two sections used here. The number of studies into
rough and scaly. Vegetative buds ovoid or conical, the phylogeny of the genus, based on DNA sequence
resinous or not resinous, often completely covered data and following cladistic methods, is still very lim-
by subterminal leaves. Leaves spirally and usually ited. Only one (Sigurgeirsson & Szmidt, 1993) included
radially arranged, firmly attached to pulvini, lin- a wide sampling (31) of species, but they analyzed only
ear, more or less equifacial (quadrangular in cross molecular data from chloroplast (cpDNA). The results
section, amphistomatic) or dorsiventral (flattened, confirm some relationships based on morphology, but
epistomatic); apex acute or acuminate (rarely ob- not others. More such work, with comprehensive sam-
tuse). Pollen cones small, 1–2(–3) cm long, ovoid pling of all species and analysing nucleotide sequences
oblong, solitary, but crowded, from axillary buds on of several genes, is needed before we can amend or
the previous year’s shoots, reddish when immature, replace Schmidt’s classification given below.
yellow when ripe; microsporophylls with two pollen
sacs containing bisaccate pollen. Seed cones subter- Genus Picea A. Dietr.
minal from long shoots, erect when immature, pen- (Type: P. abies)
dulous at maturity, soon falling or semi-persistent. Sect. Picea
Bracts only conspicuous at pollination, not growing Subsect. Picea
with the seed scales. Seed scales spirally arranged on Series Picea
a central rachis, imbricate, persistent, spreading at Species: P. abies (type of sect.,
maturity. Seeds held in a shallow cup covering one subsect. and ser.), P. obovata,
half of the seed; this membrane is continued in a P. ×fennica, P. koraiensis, P. koya­
well developed seed wing, together these are easily mae, P. asperata, P. aurantiaca,
detached from the ripe seed. Seedlings with 4–15 P. retroflexa, P. chihuahuana,
cotyledons. P. martinezii, P. crassifolia, P. mey­
eri, P. schrenkiana, P. neoveitchii,
38 species. P. torano, P. alcoquiana, P. maxi­
mowiczii, P. morrisonicola,
P. wilsonii, P. smithiana, P. glauca
Distribution
Series Rubentes Bobrov
North America: from Alaska to Newfoundland, Species: P. mariana, P. rubens
becoming disjunct southward to Mexico. Eurasia: (type), P. glehnii, P. orientalis
from the Alps and Scandinavia to Kamchatka, Sa- Subsect. Omorikae E. Murray (as “Omori­
khalin and Japan; disjunct in Caucasus/ NE Turkey; ka”)
Tian Shan; Sino-Himalayan mountain system to NE Species: P. omorika (type),
China; Taiwan. P. brachytyla, P. farreri, P. spinu­
losa, P. breweriana
 Sect. Casicta Mayr 1b. Seed cones smaller, or if longer than 10 cm less
(Type: P. jezoensis) than 5 cm wide (measured with opened seed
Subsect. Sitchenses E. Murray scales), with smaller seed scales 4
Species: P. sitchensis (type), 2a. Leaves very rigid, curved, 1.8–2.5 mm wide,
P. jezoensis, P. likiangensis, spreading radially or assurgent. Young shoots
P. linzhiensis, P. purpurea thick, very firm 3
Subsect. Pungentes E. Murray 2b. Leaves flexible, nearly straight, ca. 1mm wide,
Species: P. pungens (type), not assurgent. Young shoots thin and flexible
P. engelmannii, P. ×lutzii P. smithiana
3a. Vegetative buds large (8–12 × 4–8 mm), ovoid
552 or ovoid-oblong, smooth, shining chestnut-
Key to the sections, subsections and series of
brown; leaves strongly curved and assurgent on
Picea
thick branchlets P. torano
1a. Seed scales of cones thin and flexible, more or 3b. Vegetative buds smaller (5–6 × 3.5–4 mm),
less papery, sometimes coriaceous, usually un- conical or ovoid-conical, not smooth and shin-
dulate, with erose margins, loosely imbricate ing, slightly pubescent; leaves not assurgent, on
before ripening of the cone Sect. Casicta 2 firm but less thick branchlets P. neoveitchii
1b. Seed scales of cones rigid, more or less thin 4a. Seed cones small (3–6 × 1.5–2.5 cm), narrowly
woody, smooth or undulate, with entire or den- tapering at both ends, light red-brown when
ticulate margins, closely imbricate before rip- ripe; seed scales thin, convex, longer than wide,
ening of the cone Sect. Picea 3 with rounded or truncate apex; upper margin
2a. Leaves quadrangular in cross-section, nearly not erose. Leaves glaucous P. glauca
equifacial or bilateral, amphistomatic, glaucous 4b. Seed cones and seed scales different. Leaves
green or glaucous on all sides green or glaucous green 5
Subsect. Pungentes 5a. Seed cones relatively small (not longer than 8
2b. Leaves more or less flattened, not equifacial, cm), with obtuse apex; seed scales small (max.
epistomatic or epi-amphistomatic (with more 2 × 1.6 cm), with usually rounded, entire upper
lines of stomata on the dorsal side of the leaf), margins 6
differently coloured on two sides 5b. Seed cones usually larger (but variable accord-
Subsect. Sitchenses ing to altitude and latitude in several species!),
3a. Leaves epistomatic, usually distinctly flattened, seed scales variable 11
sometimes transversely rhombic in cross-sec- 6a. Leaves very dense above the shoot, curved 7
tion Subsect. Omorikae 6b. Leaves more remote, leaving the shoot visible
3b. Leaves amphistomatic, quadrangular (equi- from above, usually straight 8
facial) or rhombic (bilateral, keeled) in cross- 7a. Leaves pressed forward above the shoot, the
section Subsect. Picea 4 apices curved towards it. Young shoots nearly
4a. Young shoots (densely) pubescent; leaves not white; buds small (3–4 mm), slightly resinous
wider than 1(–1.5) mm, not longer than 1.5 cm. P. morrisonicola
Seed cones usually small Ser. Rubentes 7b. Leaves strongly assurgent, the apices curved
4b. Young shoots (mostly) glabrous; leaves wider upward. Young shoots yellowish to orange-
than 1 mm (usually more than 1.5 mm), if ca. 1 brown; buds larger, very resinous P. koyamae
mm wide much longer than 1.5 cm. Seed cones 8a. Young shoots (branchlets) pale buff-grey; buds
usually large Ser. Picea not resinous P. wilsonii
8b. Young shoots yellowish brown to orange-
brown; buds usually slightly resinous 9
Key to the species (and some subspecies and
9a. Leaves small (8–13 × 1–1.4 mm), spreading and/
varieties, but excluding hybrids) of Picea
or appressed forward above the shoot
Sect. Picea, Subsect. Picea, Ser. Picea
P. maximowiczii
1a. Seed cones large, broad (8–15(–18) × 4–7 cm), 9b. Leaves larger, spreading radially 10
with very large, convex seed scales (2–3 × 1.5–3
cm), with incurved, rounded margins 2
10a. Young shoots (pale) yellowish brown or pale 19b. Buds not resinous (or only slightly so); bud
red-brown; leaves with obtuse or acute apex scales appressed 20
P. koraiensis 20a. Bark grey, inner bark orange P. schrenkiana
10b. Young shoots bright orange or red-brown; 20b. Bark orange-brown to red-brown P. crassifolia
leaves with acute apex P. obovata 21a. Young shoots pale yellowish brown, entirely
11a. Seed scales with a narrowly elongated, often glabrous; buds slightly resinous 22
emarginate apex (in P. abies var. abies cones 21b. Young shoots orange or orange-brown, usually
with weakly elongated or even obtuse-rounded variously pubescent; buds resinous 23
seed scales occur, but these types are less com- 22a. Leaves with 3–5 lines of stomata on each face;
mon) 12 upper margin of seed scales entire. Resin canals
11b. Seed scales not elongated at apex, but obtuse, in leaves usually present P. martinezii 553
rounded or truncate 17 22b. Leaves with 4–10 lines of stomata on each face;
12a. Seed cones small (4–7.5 cm long); apex of seed upper margin of seed scales denticulate. Resin
scales entire, reflexed P. alcoquiana var. reflexa canals in leaves absent P. chihuahuana
12b. Seed cones usually longer than 6 cm; apex of 23a. Young shoots sparsely pubescent; buds gla-
seed scales emarginate, (slightly) incurved 13 brous at base. Seed scales not spreading wide or
13a. Seed cones 6–10(–12) cm long. Leaves dense, reflexed in ripe cones 24
curved forward; stomata in 1–3 lines on each 23b. Young shoots often ferruginous pubescent;
ventral face, in 3–6 lines on each dorsal face buds pubescent at base. Seed scales spreading
P. alcoquiana var. alcoquiana wide or slightly reflexed in ripe cones
13b. Seed cones 12–16 cm long. Leaves radially P. retroflexa
spreading; lines of stomata equally numerous 24a. Seed scales large (2–2.4 × 1.5–1.8 cm), with
on all sides of leaves 14 nearly rounded apex P. aurantiaca
14a. Seed scales rhombic-oblong; apex strongly 24b. Seed scales smaller (1.2–2 × 0.8–1.6 cm), with
elongated 15 obtuse or rounded apex
14b. Seed scales angular-obovate (obtrullate) to P. asperata var. asperata
more or less rhomboid; apex slightly elongated
16
Key to the species (and some subspecies and
15a. Leaves (especially young leaves) glaucous
varieties, but excluding hybrids) of Picea
green; branches not pendulous
Sect. Picea, Subsect. Picea, Ser. Rubentes
P. asperata var. notabilis
15b. Leaves (dark) green; branches spreading or 1a. Seed cones narrowly cylindrical, tapering to-
pendulous P. abies var. acuminata wards apex, usually 5–9 cm long. Leaves less
16a. Young leaves glaucous green than 1 cm long, dark glossy green on all sides,
P. asperata var. ponderosa some white lines of stomata on ventral side,
16b. Young leaves green P. abies var. abies more lines on the dorsal side. Basal scales of
17a. Seed cones narrowly tapering towards apex; buds obtuse-triangular P. orientalis
seed scales rounded or truncate at apex. Young 1b. Seed cones ovoid, ovoid-oblong or broad-­
shoots pubescent P. alcoquiana var. acicularis cylindrical, with obtuse or truncate apex. Leaves
17b. Seed cones obtuse at apex, more cylindrical; longer than 1 cm (if shorter, cones smaller than
seed scales obtuse or rounded at apex. Young 4 × 2.8 cm), with a different colour. Basal scales
shoots glabrous or pubescent 18 of buds acute-cuspidate 2
18a. Seed cones narrowly cylindrical, usually 7–11 2a. Seed scales undulate, with undulate or emar-
× 2.5–3.5 cm, of very regular shape; seed scales ginate, erose-denticulate upper margin; cones
obovate, broad, with very regularly rounded 3.5–8.5 cm long, often with a truncate apex
apex 19 P. glehnii
18b. Seed cones broadly cylindrical, 8–14 × 3–5 cm, 2b. Seed scales not undulate, abaxial surface rough
of less regular shape; seed scales with more or or smooth, upper margin not emarginate;
less rounded or obtuse apex 21 cones smaller (max. 6 cm long), with an obtuse
19a. Buds resinous; bud scales recurved near bud apex 3
apex P. meyeri
 3a. Seed cones very small (1.5–4 × 1.5–2.8 cm), of- mm long (nearly half length of seed scales)
ten numerous in the top of the tree, persisting P. sitchensis
several years. Leaves usually 0.8–1.2 cm long, 2b. Leaves broader (1.5–2 mm), obtriangular in
dark green or glaucous green above P. mariana cross-section, acutish or mucronate. Bracts
3b. Seed cones usually larger (2.5–6 × 1.8–3.5 cm), 4–5 mm long (less than a third length of seed
soon deciduous after shedding seeds. Leaves scales) P. jezoensis subsp. jezoensis
usually 1–1.5 cm long, shiny light green above 3a. Seed cones small (2.5–5 × 1.7–3 cm), purple,
P. rubens violet or crimson when immature; seed scales
rhombic, with incurved, papery apex
P. purpurea
Key to the species (and some subspecies and
554 3b. Seed cones larger or with different seed scales 4
varieties, but excluding hybrids) of Picea
4a. Young shoots glabrous, reddish brown in the
Sect. Picea, Subsect. Omorikae
second year; buds not resinous; leaves dis-
1a. Leaves transversely rhombic in cross-section, tinctly flattened, epistomatic. Seed cones with
acute and pungent at apex, up to 3.5 cm long, nearly obovate, flat seed scales
radially spreading P. spinulosa P. jezoensis subsp. hondoensis
1b. Leaves distinctly flattened, obtuse or acutish, 4b. Young shoots lighter, yellowish, or pubescent;
not pungent, pressed against the shoot above, buds (slightly) resinous; leaves variable. Seed
pectinate below 2 cones mostly with rhombic or less often with
2a. Seed cones small (5–8 × 2–3 cm); seed scales angular-obovate (obtrullate) seed scales 5
suborbicular, remaining imbricate. Young 5a. Young shoots brown pubescent, hairs glandu-
shoots distinctly pubescent P. omorika lar; leaves (almost) epistomatic P. linzhiensis
2b. Seed cones larger; seed scales of different shape, 5b. Young shoots glabrous or slightly pubescent,
spreading after ripening of the cone. Young hairs not glandular; leaves amphistomatic
shoots glabrous to pubescent 3 P. likiangensis
3a. Seed scales obovate, convex, with a rounded,
more or less incurved upper margin 4
Key to the species (and some subspecies and
3b. Seed scales obovate-oblong or rhombic, with
varieties, but excluding hybrids) of Picea
(usually) recurved and sometimes elongated
Sect. Casicta, Subsect. Pungentes
upper margin P. brachytyla
4a. Leaves long and wide (15–35 × 1.5–2 mm), 1a. Seed cones of medium size (5–8 × 3–4.5 cm);
curved and spreading. Branches of second or- seed scales obtrullate or rhombic; apex usually
der extremely long pendulous. Seed cones ses- narrowly elongated. Leaves more or less assur-
sile; seed scales regular, with rounded margins gent, rigid; apex very acute, pungent P. pungens
and much resin P. breweriana 1b. Seed cones usually small (3–6 × 2–2.5 cm); seed
4b. Leaves smaller (15–25 × 1–1.1 mm), straight. scales obovate-obtrullate; apex not narrowly
Branches shorter and less pendulous. Seed elongated. Leaves directed forward, flexible;
cones with an oblique peduncle; seed scales apex acute but not pungent 2
obovate, but more irregularly shaped P. farreri 2a. Leaves 1.5–2 mm wide
P. engelmannii subsp. engelmannii
2b. Leaves 1–1.2 mm wide
Key to the species (and some subspecies and
P. engelmannii subsp. mexicana
varieties, but excluding hybrids) of Picea
Sect. Casicta, Subsect. Sitchenses
1a. Immature seed cones yellowish green or green, Picea abies (L.) H. Karst., Deutsche Fl.: 324. 1881.
ripening to yellowish brown or (pale) reddish
rown 2
Etymology
1b. Immature seed cones red or purple, ripening to
purplish brown or (dark) red-brown 3 The species epithet ‘abies’, given by Linnaeus under
2a. Leaves thin (ca. 1 mm), nearly quadrangular in Pinus, is the classical name for fir (Abies), not for
cross-section, acute and pungent. Bracts 5–8 spruce (Picea), and has led to much confusion.
 between the two species ‘fuzzy’ which led to the pos-
Vernacular names
tulated hybrid taxon Picea ×fennica (Regel) Komar-
Norway spruce; Gemeine Fichte (German); Epicéa ov. Other botanists have included these populations
commun (French); Jel europeiskaya (Russian) with Picea abies subsp. obovata (Ledeb.) Hultén, a
view which would either extend the range of Picea
abies across Siberia or that of P. obovata into Scan-
Description
dinavia, depending on the taxonomic rank one
Trees to 40–50(–60) m tall, d.b.h. to 1–1.5 m; trunk prefers. The variability in Central Europe has given
monopodial, straight; bark becoming grey at low- rise to the recognition of numerous additional taxa,
er part of trunk, rough, scaly, breaking into small at ranks from species to forma. Only two varieties
plates. Branches of first order slender, spreading are recognized here: var. abies and var. acuminata. 555
horizontally or curved downward; branches of sec- There are several distinct ecotypes within this spe-
ond order highly variable, spreading horizontally cies, which are not named taxonomically.
(German: Plattenfichte) or extremely pendulous
(Kammfichte); crown (broad) pyramidal or narrow-
Distribution
ly conical. Branchlets slender, bright orange or red-
brown, becoming grey, smooth, ridged and grooved, Central (on mountains), N and E Europe, eastward
(sparsely) pubescent or glabrous; pulvini 1 mm, ob- to the Ural Mts., where the species merges with
lique. Vegetative buds ovoid conical, 4–5 × 3–4 mm, P. obovata.
not or slightly resinous; bud scales triangular-ob- TDWG codes: 10 FIN NOR SWE 11 AUT-AU AUT-
tuse, light brown or reddish brown, persisting sev- LI CZE-CZ CZE-SK GER HUN POL SWI 12 FRA-FR
eral years. Leaves spreading radially around shoot, 13 ALB BUL GRC ITA-IT ROM TUE YUG-BH YUG-CR
directed forward, parting below, 0.8–2(–3) cm long, YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 BLR
1–1.8 mm wide, linear, straight or curved, quadran- BLT-ES BLT-LA BLT-LI RUG RUE RUN RUW UKR-MO
gular in cross-section, acute at apex; amphistomatic, UKR-UK
with 2–4 lines of stomata on each face; leaf colour
(dark) green. Pollen cones near end of shoot, 1–1.5
Ecology
cm long, yellow. Seed cones terminal, sessile, cy-
lindrical, rarely ovoid-oblong, (2.5–)6–15(–20) cm Picea abies is widespread and dominant in boreal
long, 1.5–4(–5) cm wide with opened scales; colour conifer forests of N and NE Europe, where it replac-
(immature) green or red, maturing to orange-brown es Pinus sylvestris on wetter sites because Picea abies
or (light) red-brown. Seed scales highly variable, can avoid the water table with a very shallow root
ovoid-oblong to rhomboid-oblong or rhombic, 1.5–3 system. The natural distribution shows continental
× 1–2 cm at mid-cone; surface smooth, shining, gla- tendencies but in the western mountains of Cen-
brous; upper margin variable, obtuse or acuminate, tral Europe an ecotype has evolved that is adapted
with retuse apex, entire or erose-denticulate; base to sub-Atlantic weather conditions with heavy ‘wet’
cuneate. Bracts rudimentary, ligulate, 2–3 mm, en- snowfall in early winter. Its inability to compete with
tirely included. Seeds ovoid-oblong, 2–5 mm, dark more shade tolerant Abies alba and Fagus sylvatica
brown or blackish brown; seed wings ovate-oblong as well as historical factors have limited its natural
or cuneate, (6–)10–20 mm long, light brown. expansion into W Europe. In the Alps Picea abies
occupies the montane to subalpine zones (depend-
ent on local climate) especially on moist sites and
Taxonomic notes
in cold air pockets. Although it can occur on most
Populations in northern Scandinavia and especially substrates, acidic soils are most common and wide-
in NE Russia differ from the otherwise very variable spread as is testified by the undergrowth, if present,
species elsewhere in having smaller cones with more of ericaceous shrubs and sub-shrubs. Commonly
or less rhombic seed scales with obtuse, often ser- growing with Picea abies in the boreal forests are
rulate margins and more pubescent shoots. These Betula sp. and Populus tremula, with willows (Salix)
characters are intermediate between P. abies and alongside streams and lakes. In the Alps Picea abies
P. obovata and in some populations even seem to occurs with Larix decidua, Pinus cembra, and P. syl­
merge into those of the latter, making the ­boundary vestris or P. nigra, if not in pure stands. In E Europe,
 Picea abies is a constituent of mixed conifer-broad- TDWG codes: 10 FIN NOR SWE 11 AUT-AU AUT-
leaved woodland from the Bialowiecza Forest in the LI CZE-CZ CZE-SK GER HUN POL SWI 12 FRA-FR
north to the valleys of the eastern Alps and the Car- 13 ALB BUL GRC ITA-IT ROM TUE YUG-BH YUG-CR
patians. YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 BLR
BLT-ES BLT-KA BLT-LA BLT-LI RUC RUE RUN RUW
UKR-MO UKR-UK
Uses
Norway spruce is an important timber tree in
Conservation
­Europe, where outside the boreal forest zone most
commercial timber is now harvested from planta- IUCN: LC
556 tions or from managed forests in which other trees
are suppressed. Forestry has expanded the range of Picea abies (L.) H. Karst. var. acuminata (Beck)
this species considerably further west. The wood Dallim. & A. B. Jacks., Handb. Conif., ed. 2: 390.
is used for pulpwood as well as construction, fur- 1931. Picea excelsa (Lam.) Link var. acuminata
niture (most of the popular ‘pine’ furniture is made Beck, Ann. K. K. Naturhist. Hofmus. 2: 61. 1887.
with wood from Norway spruce), and special uses Type: Illustration in M. Kienitz, Über Formen
like the sound boards of pianos and the bodies of und Abarten heimischer Walbäume, t. 3, f. 5a. 1879
guitars and violins. The famous Stradivarius vio- (holotype).
lins were made with wood of Norway spruce from
the Alps. In Europe this species is the most popu-
Description
lar Christmas tree, a tradition that actually started
in Germany, with the extensive afforestation begin- Seed cones generally large, 12–16 cm long; seed
ning in the 18th century. Norway spruce is not much scales rhombic, with narrowly elongated, acuminate
planted as an amenity tree, but in horticulture more and incurved, erose-denticulate apex.
than 200 cultivars have been selected, with differ-
ent habits including ‘weeping’, prostrate and dwarf
Distribution
forms, red, white or yellow flushing leaf forms, and
(other) monstrosities. Europe: Jura, Alps, Carpathian Mts., Sweden?
TDWG codes: 11 AUT-AU CZE-CZ CZE-SK POL
2 varieties are recognized: ROM SWI 12 FRA-FR 13 ITA-IT 14 UKR-UK

Picea abies (L.) H. Karst. var. abies. Pinus abies L.,

Conservation
Sp. Pl. 2: 1002. 1753. Type: Illustration “Picea” in
Camerarius, Pl. Epit.: 47. 1586 (lectotype). IUCN: LC

Abies alpestris Brügger, Jahresber. Naturf. Ges.

Graubündens, ser. 2, 29: 167. 1886; Picea alpestris Picea alcoquiana (Veitch ex Lindl.) Carrière, Traité
(Brügger) Stein, Gartenfl. 37: 346. 1887; Picea abies Gén. Conif., ed. 2, 1: 343. 1867. Pl. 23
(L.) H. Karst. var. alpestris (Brügger) P. A. Schmidt,
Haussknechtia 4: 38. 1988.
Etymology
This species was named after Rutherford Alcock,
Description
at the time H. M. Minister at the Court of Jeddo in
Seed cones variable in size, (2.5–)6–15(–20) cm long; ­Japan.
seed scales ovoid-oblong to rhomboid-oblong; apex
variable but not narrowly elongated.
Vernacular names
Alcock’s spruce; Matsuhada, Ira-momi (Japanese)
Distribution
Central, N and E Europe, eastward to the Ural
Mountains.
 are frequent. The forests on these mountains are
Description
mixed coniferous, with Picea jezoensis subsp. hon­
Trees to 30–35 m tall, d.b.h. to 1 m; trunk mono­podial, doensis as the most common of the spruces, Tsuga
straight; bark of trunk dark grey, breaking into ir- diversifolia and Larix kaempferi, both also common,
regular plates. Branches of first order long, slender, Pinus parviflora and Abies veitchii in some areas,
spreading horizontally, but lower ones downward; A. mariesii usually at higher elevations, and broad-
branches of second order spreading or slightly as- leaved trees, e.g. Betula ermanii, B. grossa, Sorbus
surgent; crown (broad) pyramidal or conical, in old commixta, Quercus mongolica var. grosseserrata, Al­
trees open. Branchlets slender, firm, yellowish brown nus hirsuta var. sibirica, and Prunus maximowiczii.
or pale orange-brown, smooth, shining, ridged and
grooved, glabrous or more or less pubescent; pulvini 557
Uses
small, 0.5 mm, at 50–80° from shoot axis. Vegetative
buds ovoid-conical or subglobose, 3–5 mm long, res- Alcock spruce is a timber tree of minor importance
inous; bud scales triangular, brown, persisting sever- due to its scarcity, but it has undoubtedly been
al years. Leaves curved forward above shoot, parted logged with other spruces (P. jezoensis subsp. hon­
and spreading below, 0.8–1.5(–2.5) cm long, 1–1.5 doensis) and conifers. Much of its wood is processed
mm wide, linear, often curved, quadrangular; apex to pulp for the paper industry, but more specialized
acute, pungent in young trees; stomata on all sides, uses are furniture making and (in Japan) musical
in 1–3 lines on each ventral face, in 3–6 lines on each instruments. This species has been introduced to
dorsal face; leaf colour dark blue green, with bluish Europe and the USA but remains uncommon there
white stomatal lines. Pollen cones axillary, clustered, and mostly restricted to arboreta and some large
1–1.5 cm long, rose-red, later yellow. Seed cones ter- parks of private estates, where it may still be known
minal, erect at first, soon pendant, sessile or very under the later name Picea bicolor.
short pedunculate, ovoid-oblong, (4–)6–10(–15) cm
long, (2.5–)3–5(–5.5) cm wide with opened scales, 3 varieties are recognized:
violet when immature, ripening to reddish brown
or (pale) brown. Seed scales obovate-rhombic to Picea alcoquiana (Veitch ex Lindl.) Carrière var.
oblong-spathulate, opening wide, 1.5–2.5 × 0.8–1.5 alcoquiana. Abies alcoquiana Veitch ex Lindl.,
cm at mid-cone, thin and flexible, undulate; surface Gard. Chron. 1861: 23. 1861 [“alcockiana”]. Type:
smooth, finely striated, glabrous, upper margin un- Japan, Honshu, Fuji-yama, coll. J. G. Veitch (type
dulate, serrulate, notched at apex and often reflexed, not designated).
or entire and rounded to truncate, base narrow
cuneate, dark brown. Bracts rudimentary, ligulate, Abies bicolor Maxim., Bull. Acad. Imp. Sci. Saint-
2–3 mm, entirely included. Seeds ovoid-oblong, 2–3 Pétersbourg 10: 488. 1866; Picea bicolor (Maxim.)
mm long, blackish brown; seed wings ovate-oblong, Mayr, Monogr. Abiet. Japan. Reich.: 49. 1890.
13–18 mm long, light yellowish brown.
Description
Distribution
New shoots glabrous; leaves 0.8–1.5(–2) cm long,
Japan: central Honshu (mainly in Kanto Mts. and slightly curved forward. Seed cones (5–)6–10(–12)
Chubu Mts.). cm long; seed scales thin, flexible, spreading wide;
TDWG codes: 38 JAP-HN upper margin undulate, notched at apex, serrulate.

Ecology Distribution
Picea alcoquiana is a scattered mountain species, oc- Japan: central Honshu.
curring at elevations from 700 m to 2180 m (var. ac­ TDWG codes: 38 JAP-HN
icularis between 1200 m and 1950 m) a.s.l. The soils
are of volcanic origin and podzolic. The climate is
Conservation
cool, with cold, snowy winters, and wet (annual pre-
cipitation 1000 mm to 2500 mm), while typhoons IUCN: LC
 Picea alcoquiana (Veitch ex Lindl.) Carrière var. Picea asperata Mast., J. Linn. Soc., Bot. 37: 419.
acicularis (Maxim. ex Beissn.) Fitschen, in Beissn- 1906.
er, Handb. Nadelholzk., ed. 3: 258. 1930. Picea acicu­
laris Maxim. ex Beissn., Handb. Nadelholzk.: 380.
Etymology
1891; Picea bicolor (Maxim.) Mayr var. acicularis
(Maxim. ex Beissn.) Shiras., Bot. Mag. (Tokyo) 27: The species epithet (Latin asperatis = roughness)
130. 1913. Type: Japan, coll. C. J. Maximowicz (type perhaps refers to the foliage branches.
not designated).
Vernacular names
Picea shirasawae Hayashi, Ill. Useful Trees (Forest
558 Trees): sub f. 43. 1969. Dragon spruce; yunshan (Chinese)

Description Description
New shoots pubescent; leaves strongly curved, glau- Trees to 30–45 m tall, d.b.h. to 1–1.5 m; trunk mono­
cous, 1.3–2.5 cm long. Seed cones 6–15 cm long with podial, straight; bark orange brown, turning grey
a narrowed apex; seed scales with a rounded or brown, in large trees rough and scaly. Branches of
truncate, entire or denticulate upper margin. first order relatively short, spreading horizontally,
upturned at ends; branches of second order spread-
ing or slightly assurgent; crown (narrowly) coni-
Distribution
cal, gradually tapering towards the top. Branchlets
Japan: Honshu (Yatsugadake Mts.). slender, firm, shining yellowish brown or orange,
TDWG codes: 38 JAP-HN turning grey with age; surface smooth, prominently
ridged and deeply grooved, glabrous, or variously,
but not densely, pubescent; pulvini rigid, 2 × 1 mm,
Conservation
erect or recurved. Vegetative buds ovoid-conical
IUCN: VU (D2) or conical, acute, 6–12 × 5–8 mm, resinous; bud
scales triangular acute, keeled, appressed, or the api-
Picea alcoquiana (Veitch ex Lindl.) Carrière var. cal scales slightly recurved, yellowish brown with
reflexa (Shiras.) Fitschen, in Beissner, Handb. reddish margins, persisting several years. Leaves
Nadelholzk., ed. 3: 258. 1930. Picea bicolor (Maxim.) spreading radially, forward or assurgent above, part-
Mayr var. reflexa Shiras., Bot. Mag. (Tokyo) 27: ed and spreading laterally below, 1–2(–2.5) cm long,
(129), t. 2, f. 9–12. 1913. Type: Illustration in Bot. 1–1.8 mm wide (widest at base), linear, curved or
Mag. (Tokyo) 27, t. 2, f. 9–12. 1913. (holotype). straight, quadrangular-rhomboid in cross-section,
acuminate and very pungent at apex; amphistomat-
ic, with 3–4 lines of stomata on each face; colour of
Description
young leaves glaucous green, later dull green. Pollen
New shoots pubescent; leaves curved forward, 0.8– cones near end of shoots, axillary, 1–1.5 cm long, red-
1.3 cm long. Seed cones 4–7.5 cm long; seed scales dish, then yellow. Seed cones terminal, erect at first,
entire, apically narrowed and reflexed. pendulous at maturity, short pedunculate or sessile,
± cylindrical, tapering at both ends (fusiform), ob-
tuse at apex, (5–)6–12(–15) cm long, (2.5–)3–4(–4.5)
Distribution
cm wide with opened scales; colour (immature)
Japan: Honshu (Akaishi Range). purplish green or purple, ripening to (dark) brown
TDWG codes: 38 JAP-HN or reddish brown. Seed scales obovate or rhombic-
oblong, 1.2–2 × 0.8–1.6 cm at mid-cone; surface
striated to nearly smooth, glabrous; upper margin
Conservation
obtuse, rounded or truncate to emarginate, usually
IUCN: VU (D2) slightly incurved; base cuneate. Bracts rudimentary,
ligulate, 2–3 mm, entirely included. Seeds ovoid,
with acute apex, 2–4 × 1.5–2.8 mm, dark brown or
 559

5 6
1

7
3

Pl ate 2 3. Picea alcoquiana. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone (var. alcoquiana). 4. Seed
scale (var. alcoquiana). 5. Seed cone (var. acicularis). 6. Seed scale (var. acicularis). 7. Seed cone (var. reflexa).
8. Leaf. 9. Seeds.
 grey-brown; seed wings ovate-oblong, 8–12 × 5–6 China’s rapidly growing economy. The species and
mm, yellowish brown, transparent. its varieties are in cultivation as amenity trees and in
arboreta and pineta in Europe and the USA, mostly
still based on seed collections by Ernest Wilson,
Taxonomic notes
Joseph Rock and some other early 20th century plant
Picea asperata is a variable Chinese spruce, not un- hunters who travelled widely in southern Gansu and
like Picea abies in Europe, and it might be better western Sichuan.
taxonomic practice to recognize this as such and
not to name varieties as they seem to appear in the 3 varieties are recognized:
limited numbers of specimens (both in herbaria and
560 grown from seed in arboreta) available for study. Picea asperata Mast. var. asperata. Type: China:
Population based studies in the field on P. abies have Sichuan, [“Western China”], E. H. Wilson E. H. 3025
revealed regional forms or ‘races’, some related to (holotype BM).
environmental factors, as well as much introgres-
sion. If such detailed studies were available for P. as­
Description
perata, similar patterns would be likely. As some of
the varieties have been traditionally recognized and New shoots glabrous or variously pubescent; bud
since the above suggested studies are not available, I scales mostly appressed. Seed cones (5–)6–12 cm
have tentatively retained them here. long, (2.5–)3–4 cm wide when opened; seed scales
obovate-oblong, with obtuse, rounded or truncate
upper margin.
Distribution
China: Gansu, Ningxia (Helan Shan), E Qinghai,
Distribution
SW Shaanxi, Sichuan.
TDWG codes: 36 CHC-SC CHI-NX CHN-GS CHN-SA China: Gansu, Ningxia (Helan Shan), E Qinghai,
CHQ SW Shaanxi, Sichuan.
TDWG codes: 36 CHC-SC CHI-NX CHN-GS CHN-SA
CHQ
Ecology
Picea asperata occurs in the high mountains of
Conservation
W central China, at elevations between 1500 m and
3800 m a.s.l., usually above 2400 m in Sichuan. The IUCN: LC
soils are grey-brown mountain podzols. The climate
is continental, subalpine, with cold winters and dry Picea asperata Mast. var. notabilis Rehd. &
summers (annual precipitation less than 500 mm). E. H. Wilson, in Sargent, Pl. Wilson. 2: 23. 1914.
It forms mostly pure forests on N-facing slopes, or Picea notabilis (Rehd. & E. H. Wilson) Lacass., Trav.
mixtures with other species of Picea, in the south of Lab. Forest. Toulouse T. 1 (3, 1): 180. 1932. Type:
Gansu it may be mixed with Abies nephrolepis. Bet­ China: Sichuan, Guan Xian, Banlan Shan, [“West
ula albo-sinensis is the most common broad-leaved of Kuan Hsien, Panlan Shan”], E. H. Wilson 2068
associate. (holotype A).

Uses Description
Picea asperata is an important timber tree in China. Young shoots glabrous, bright orange brown; bud
The wood is mainly used for pulpwood and to a less- scales loosely imbricate, apically reflexed, yellowish
er extent for construction. Old growth stands of this brown. Seed scales rhombic oblong, emarginate.
potentially large spruce have been reduced to less
accessible mountain slopes and valleys and planta-
Distribution
tion forestry has begun to replace the natural stands
as a resource for spruce timber, but as yet on a scale China, W Sichuan.
that is incapable of meeting growing demands in TDWG codes: 36 CHC-SC
 pendulous; crown narrowly conical in large trees,
Conservation
open. Branchlets thick, firm, rigid, orange or yel-
IUCN: LC lowish orange, slightly pruinose, becoming grey with
age; surface smooth, deeply grooved, with flat ridg-
Picea asperata Mast. var. ponderosa Rehd. & es, young shoots sparsely pubescent, soon glabrous;
E. H. Wilson, in Sargent, Pl. Wilson. 2: 23. 1914. pulvini well developed, 1–2 mm, at right angle from
Picea ponderosa (Rehd. & E. H. Wilson) Lacass., shoot. Vegetative buds broad ovoid-conical, lateral
Trav. Lab. Forest. Toulouse T. 1 (3, 1): 203. 1932. buds subglobose, 4–6 × 4–6 mm, covered with pul-
Type: China: Sichuan, Guan Xian, Banlan Shan vini at base, hidden by leaves, resinous; bud scales
[“West of Kuan Hsien, Panlan Shan”], Panlan Shan, triangular, obtuse or acutish, yellowish brown, with
E. H. Wilson 4068 (holotype A). red-brown, erose margins, persisting several years. 561
Leaves all around the shoot, curved or directed for-
ward, on shaded shoots parted below, more or less
Description
pectinate, 0.8–1.8 cm long, 1.2–2.3 mm wide, linear,
Seed cones usually large, 12–15 × 4–4.5 cm; seed straight or curved, quadrangular-rhomboid in cross-
scales coriaceous, rigid, with slightly elongated, section, with ribs on two opposite sides, acute-mucro­
curved and usually emarginate apex. nate, pungent; amphistomatic, stomata in 4 bands of
3–6 lines; young leaves bluish green, later grey-green.
Pollen cones near end of shoot, axillary, 1.5–2 cm
Distribution
long, reddish, then yellow with pollen. Seed cones
China: W Sichuan. terminal, erect at first, pendulous at maturity, ses-
TDWG codes: 36 CHC-SC sile or short pedunculate, cylindrical-oblong, obtuse
at apex, 10–12 cm long, 4–4.5 cm wide with opened
scales; immature cones bright red, maturing to red-
Conservation
dish brown; ripe cones shining chestnut brown or
IUCN: LC dull brown. Seed scales rhombic-ovoid, with nearly
rounded apex, 2–2.4 × 1.5–1.8 cm at mid-cone; sur-
face striated, shining, glabrous; upper margin slightly
Picea aurantiaca Mast., J. Linn. Soc., Bot. 37: 420. erose denticulate; base cuneate. Bracts rudimentary,
1906. Picea asperata Mast. var. aurantiaca (Mast.) ligulate, 2–4 mm, entirely included. Seeds ovoid-
Boom, in Den Ouden & Boom, Man. Cult. Conif.: conical, 3–4 mm long, brown; seed wings ovate ob-
253. 1965. Type: China: Sichuan, Daxue Shan, Che- long, 12–14 × 5–6 mm, transparent, yellowish brown.
to Shan, [“West of Tachien-lu”], E. H. Wilson 3029
(holotype BM).
Taxonomic notes
Masters (op. cit.), in his lecture to the Linnean So-
Etymology
ciety, separated this species from the closely related
The Latin species epithet means ‘orange’ and refers species P. asperata Mast. and P. retroflexa Mast., all
to the colour of the foliage branches. three newly described there. He already made alle-
gations about their close relationship and possible
Vernacular names intermediate forms. These taxa have been variously
treated as independent species or varieties of P. as­
Orange spruce; baipi yun shan (Chinese)
perata (Schmidt-Vogt, 1977). In Flora of China 4:
28 (1999) and in Higher Plants of China 3: 37 (Fu
Description
et al., 2000) P. aurantiaca is treated as a variety of
Trees to 20–25 m tall, d.b.h. to 0.7 m; trunk mo- P. asperata, while other accounts (e.g. Fu & Jin, 1992;
nopodial, straight; bark becoming scaly, grooved Farjon, 1990, 1998, [2001]) have maintained the
and ridged, strongly exfoliating, with irregular plates, species rank. A re-examination of relevant collec-
pale grey. Branches of first order short, spreading tions and populations seems desirable; this should
horizontally or curved down, sparse in old trees; ­include work on DNA sequences, looking for mark-
branches of second order thick, rigid, spreading, not ers which may help to distinguish species.
 Distribution Vernacular names
China: W Sichuan (Zheduo Shan W of Kangding, Sargent’s spruce; mai diao yun shan (Chinese)
from Simaqiao to Xinyulingong).
TDWG codes: 36 CHC-SC
Description
Trees to 30(–40) m tall, d.b.h. to 1–1.2 m; trunk mo-
Ecology
nopodial, straight; bark becoming rough, scaly, flak-
Picea aurantiaca is a subalpine species, occurring be- ing, dark grey, with reddish brown freshly exposed
tween 2600 m and 3800 m a.s.l. (–4000 m according bark plates. Branches of first order long, slender,
562 to Rehder & Wilson, 1914). It is mostly found on cal- spreading horizontally; branches of second order
careous soils. The climate is cold and precipitation slender, pendulous; crown broad conical, open in
varies from high (no figures recorded) in the lower old trees. Branchlets slender, flexible, leading shoots
elevations of the SE of its range to only 500–700 mm stout, creamy white at first, later light brown to
in the NW. It occurs in mixed coniferous forest, with orange-brown, sometimes pruinose; surface finely
e.g. Picea likiangensis var. rubescens, Abies squamata, ridged and grooved, glabrous or minutely pubes-
A. chensiensis, A. recurvata, Tsuga chinensis, and lo- cent; pulvini small, whitish yellow, at 45–60° to the
cally, Larix potaninii. Betula spp. are the common shoot axis. Vegetative buds ovoid conical, on leading
broad-leaved trees, while Pinus spp. occur mostly shoots 5–8 × 4 6 mm, smaller on lateral shoots, cov-
after disturbances and at the lower elevations. ered by leaves, (slightly) resinous; bud scales closely
appressed, triangular, chestnut brown, persistent.
Leaves curved forward and downward, with lower
Conservation
leaves parted, nearly pectinate, on coning shoots
The limited areas of occurrence and occupancy more radially spreading, (0.8–)1–2(–2.4) cm long,
(EOO and AOO) estimated for this species and the 1–1.5(–2) mm wide, truncate at base, linear, curved,
inferred decline from logging operations and exten- slightly flattened, keeled on both sides; apex acute or
sive deforestation in the area justify the Red List cat- mucronate; stomata on lower side only, in 2 narrow
egory Endangered. bands separated by a midrib; leaf colour dark green
IUCN: EN (B1ab+2ab) above, white bands below. Pollen cones axillary,
1–2 cm long, yellowish when shedding pollen. Seed
cones terminal, erect at first, pendulous at maturity,
Uses
short pedunculate, cylindrical-oblong or ovoid-ob-
A timber tree of which no further details of its uses long, (obliquely) tapering at base; apex obtuse, (5–)
are recorded, presumably because it is not recog- 6–10(–15) cm long, 3–4(–5) cm wide with opened
nized as distinct from Picea asperata. It must be scales colour (immature) green or purplish green,
logged with this and other spruces and put to the ripening to dark brown with a purplish band across
same uses. It was introduced in England and is still each scale. Seed scales angular-bovate to nearly
found in some arboreta in the south, growing par- rhombic, thin, rigid, 1.6–2(–2.5) × 1–1.4(–1.8) cm at
ticularly well on shallow soil over chalk. mid-cone; abaxial surface striated or wrinkled, gla-
brous; upper margin variably undulate or rounded
to truncate, usually recurved; apex emarginate or
Picea brachytyla (Franch.) E. Pritz., Bot. Jahrb. erose; base cuneate. Bracts rudimentary, ligulate,
Syst. 29: 216. 1900. 2–3 mm, entirely included. Seeds ovoid-oblong, 3 ×
2 mm, light brown; seed wings ovate-oblong, 10–14
× 5–7 mm, orange-brown.
Etymology
The species epithet (Greek: brachy = short; tylo =
Distribution
‘with lumps or projections’) refers to the short pul-
vini on the shoots. China: S Gansu, NW Hubei, S Shaanxi, W Sichuan,
NW Yunnan, SE Xizang [Tibet]; NE India: Aru-
nachal Pradesh (Assam Himalaya); N Myanmar
Description
[Burma]; Bhutan (?).
TDWG codes: 36 CHC-HU CHC-SC CHC-YN CHN- Seed cones 6–10(–12) cm long, 3–4 cm wide when
GS CHN-SA CHT 40 EHM-AP 41 MYA opened; seed scales angular-obovate, 1.6–2 cm long,
1–1.4 cm wide; upper margin variably undulate,
emarginate or erose, usually recurved.
Ecology
Picea brachytyla is a high mountain species, occur-
Distribution
ring between 1300 m and 3800 m a.s.l. The soils are
grey-brown mountain podzols. The climate is cold China: Chongqing, S Gansu, NW Hubei, S Shaanxi,
and wet, with annual precipitation from 1000 mm Sichuan, NW Yunnan, SE Xizang [Tibet]. 563
(N) to more than 2500 mm (S), where the mon- TDWG codes: 36 CHC-CQ CHC-HU CHC-SC CHC-
soon influence is stronger. It is a constituent of the YN CHN-GS CHN-SA CHT
montane coniferous forest of the eastern parts of the
Himalaya and the mountains of the SW Plateau of
Conservation
China (Wang, 1961), with Abies densa, A. forrestii,
Picea likiangensis, Pinus wallichiana, Tsuga dumosa, IUCN: LC
and Larix potaninii as major species. Taxus wallichi­
ana is commonly found as an understorey tree in Picea brachytyla (Franch.) E. Pritz. var. complanata
the Himalayan part of its range. (Mast.) W. C. Cheng ex Rehd., Man. Cult. Trees, ed.
2: 30. 1940. Picea complanata Mast., Gard. Chron.,
ser. 3, 39: 146. 1906. Type: China: Sichuan, W
Uses
Sichuan, E. H. Wilson 3031 (sheet No. 1, lectotype K,
This species is a timber tree, used for construction, designated here).
interior flooring, aircraft, machines, furniture, and
wood pulp for the paper industry. In China this Picea brachytyla (Franch.) E. Pritz. var. rhombisqua­
species has been intensively exploited, depleting mea Stapf, Bot. Mag. 148: sub t. 8969. 1923.
the natural stands, and now is cultivated for affor- Picea ascendens Patschke, Bot. Jahrb. Syst. 48: 632.
estation. In Europe and North America it is often 1913; Picea brachytyla (Franch.) E. Pritz. var. ascend­
present in larger arboreta, mainly derived from seed ens (Patschke) Silba, J. Int. Conifer Preserv. Soc. 7
collections made by European plant hunters who (1): 28. 2000.
traveled in China in the early decades of the 20th
century. It is a more attractive and shapely species
Description
than most spruces and new introductions from dif-
ferent parts of its range should be recommended for Seed cones 8–15 cm long, 3.5–5 cm wide when
horticultural uses; it is extremely hardy and tolerant opened; seed scales broadly obovate-oblong, 2–2.5
of poor soils. cm long, 1.5–1.8 cm wide at mid-cone, with usually
rounded or truncate (sometimes elongate-emargin-
2 varieties are recognized: ate) and straight or recurved upper margins.

Picea brachytyla (Franch.) E. Pritz. var. brachytyla.

Taxonomic notes
Abies brachytyla Franch., J. Bot. (Morot) 13 (8): 258.
1899. Types: China, Sichuan, P. Farges 806; Yunnan, In K there are three herbarium sheets of E. H. Wilson
A. Delavay 4129 (syntypes P). 3030 and 3031 (2 sheets) upon which Masters based
his species Picea complanata and from which the
Picea pachyclada Patschke, Bot. Jahrb. Syst. 48: 630. drawings in the protologue were made. E. H. Wilson
1913; Picea brachytyla (Franch.) E. Pritz. var. pachy­ 3031 with a mature seed cone (sheet 1) is here des-
clada (Patschke) Silba, Phytologia 68: 39. 1990. ignated as the lectotype. Stapf (1923) indicated that
E. H. Wilson 3031 belonged to his forma rhombisqua­
mea and this form is therefore treated as a synonym.
 shoots and on short branchlets, curved outward on
Distribution
both sides, 1.5–3(–3.5) cm long, 1.5–2 mm wide, lin-
China: W Sichuan, NW Yunnan; NE India: Aru- ear, flattened, convex above, weakly keeled below;
nachal Pradesh (Assam Himalaya); N Myanmar apex obtuse; epistomatic, stomata in 2 bands of 4–6
[Burma]; Bhutan (?). lines, separated by a weak midrib; leaf colour dark
TDWG codes: 36 CHC-SC CHC-YN 40 EHM-AP 41 green above, greenish white stomatal bands below.
MYA Pollen cones in leaf axils on pendulous shoots, 2–2.5
cm long, light brown to orange-brown. Seed cones
terminal, erect at first, pendulous at maturity, on
Conservation
short branches in top of tree, sessile or very short
564 Logging and deforestation have substantially re- pedunculate, cylindrical, slightly curved, oblique at
duced the extent of coniferous forests in the region base, tapered towards obtuse apex, (7–)8–12(–14)
and have had an impact especially on this variety cm long, 3–4 cm wide with opened scales; colour
with its more restricted range. (immature) green, maturing to purplish brown, rip-
IUCN: VU (A1c-d) ening to red-brown or dull brown, very resinous.
Seed scales obovate-flabellate, convex, thin but rig-
id, opening very wide when dry, 1.5–2 × 1.3–1.8 cm
Picea breweriana S. Watson, Proc. Amer. Acad. at mid-cone; surface of abaxial side smooth, slightly
Arts 20: 378. 1885. Type not designated. striated in some cones, glabrous; upper margin en-
tire, rounded or truncate, slightly incurved; base
cuneate. Bracts rudimentary, ligulate, 3 mm, entirely
Etymology
included. Seeds ovoid, 3–4 × 2–3 mm, brown; seed
This species was named after William H. Brewer wings ovate, 7–9 × 5–6 mm, orange-brown.
(1828–1910) of the California State Geological sur-
vey.
Distribution
USA: NW California, SW Oregon (Siskiyou Mts.).
Vernacular names
TDWG codes: 73 ORE 76 CAL
Brewer spruce, Weeping spruce
Ecology
Description
Picea breweriana is a mountain species of a limited
Trees to 25(–35) m tall, d.b.h. to 1 m; trunk mono­ area near the Pacific coast, at elevations between
podial, straight or curved at base; bark on trunk (1000–)1200–2300 m a.s.l. It grows in different soil
scaly, with prominent branch scars, dark reddish types, mostly associated with former glacial activ-
grey or purplish grey. Branches of first order long, ity. The climate is temperate, with usually warm, dry
slender, spreading horizontally, curved upwards, summers and cool to cold, moist winters with abun-
lower branches descending, drooping at the ends; dant snowfall. It occurs chiefly in the Transition Life
branches of second order in mature trees extremely Zone, with e.g. Abies magnifica, A. concolor, Pseudo­
long (to 2.5 m), slender, pendulous; crown broad tsuga menziesii, Pinus monticola, P. lambertiana,
conical, open and irregular in large trees. Branch- P. ponderosa, Tsuga mertensiana, Calocedrus decur­
lets slender, the leading shoots long and flexible, rens, and many shrubs (mixed conifer type). This
pink-brown or red-brown, turning grey; surface species can compete with e.g. Pseudotsuga menziesii
smooth, with ridges and shallow grooves between and Pinus monticola and is well adapted to heavy
reddish brown, 2 mm long pulvini. Vegetative buds wet snowfall (‘Kammfichte’ type of branching).
conical-oblong, with obtuse apex, 5–8 × 2.5–4.5 mm,
slightly resinous at first; bud scales triangular, ob-
Conservation
tuse, appressed, later recurved and papery, yellow-
ish brown or pale brown, persisting several years. The limited range (extent of occurrence [EOO]
Leaves spreading more or less radially on long pen- 220 × 110 km) and rarity, except for larger popula-
dulous shoots, pressed forward against the leading tions in the western Siskiyou Mountains and in the
Marble, Salmon and Trinity Mountains of northern in large trees (partly) pendulous; crown pyramidal
California, put this species potentially at risk from in young trees, in mature trees broad conical, open,
catastrophic damage. The main hazard is forest fires, irregular. Branchlets firm, slender or thick in lead-
to which the species is more vulnerable than oth- ing shoots, young shoots pale, later yellowish brown,
er conifers that occur within its range. It therefore finally grey; surface smooth, prominently ridged
retreats to N-facing slopes and rocky ridges where and deeply grooved, glabrous; pulvini strongly de-
fires are less intense or cannot reach. In the face of veloped, dense, 1–1.5 mm, on leading shoots nearly
global warming the frequency and intensity of forest erect. Vegetative buds ovoid conical, with acute apex,
fires are likely to increase, this may move P. breweri­ 4–8 × 2.5–6 mm, slightly resinous; bud scales trian-
ana within the categories of threat in future. How- gular-ovate, with ciliate-erose margins, appressed,
ever, these predictions are uncertain and therefore light orange-brown, with darker margins, persisting 565
in themselves insufficient for this species to meet the 3–4 years. Leaves on leading shoots dense, radially
criteria for a threatened category. spreading, more or less assurgent above shoot, part-
IUCN: NT ed below, 1.2–2.3(–2.8) cm long, 1–1.8 mm wide, lin-
ear, straight or curved, rigid, quadrangular-rhombic
in cross-section, acute to acuminate at apex; am-
Uses
phistomatic, 3–5 lines of stomata on each grooved
Brewer spruce is of little or no value as a timber tree. face; leaf colour grey-green or glaucous green. Pol-
It may occasionally be logged with other conifers len cones near end of shoots, axillary, 1–1.5 cm long,
and put to use as pulp wood. In horticulture, how- yellowish. Seed cones terminal, erect at first, pendu-
ever, it is considered a very desirable ornamental lous at maturity, solitary or a few together, sessile,
tree due to its long pendulous foliage branches. It is cylindrical, with obtuse apex, 10–14(–17) cm long,
widely planted especially in European arboreta and 4–5 cm wide with opened scales; colour (immature)
parks, and generally performs well, although trees bright green, ripening to shining (light) brown.
grown from seed are slow to mature to the charac- Seed scales cuneate-flabellate, convex, opening very
teristic habit; grafted trees develop faster. Few culti- wide, 2–2.5 × 1.8–2.1 cm at mid-cone; abaxial surface
vars have been selected. smooth, glabrous; upper margin entire, rounded;
base cuneate. Bracts rudimentary, ligulate, 3–4 mm,
entirely included. Seeds ovoid-cuneate, 3–6 mm
Picea chihuahuana Martínez, Anales Inst. Biol. long, dark red-brown or grey-brown; seed wings
Univ. Nac. México 13: 31. 1942. Type: Mexico: ovate-oblong, 10–15 × 5–8 mm, yellowish, tinged
Chihuahua, Bocoyna, Arroyo de los Talayotes, with red-brown.
R. Dueñas s.n. (holotype MEXU). Fig. 188
Distribution
Etymology
Mexico: SW Chihuahua, S Durango.
The species epithet refers to the Mexican State of TDWG codes: 79 MXE-CU MXE-DU
Chihuahua, from where it was first described.
Ecology
Vernacular names
Picea chihuahuana occurs in scattered relict popula-
Chihuahua spruce tions on N-facing high mountain sides, often in can-
yons, at elevations between 2150 m and 3200(–3400)
m a.s.l. It grows in poor, barren, but always moist
Description
mountain soils of alluvial origin, usually near per-
Trees to 40–45 m tall, d.b.h. to 1–1.2 m; trunk mo- manent streams, but in the Sierra Madre Oriental
nopodial, straight; bark on trunk rough and scaly, also on calcareous lithosols. The climate is cool and
breaking into numerous small plates, dark grey- moist, with annual precipitation between 800 mm
brown. Branches of first order long, slender, in and 1300 mm, mostly as summer showers, but in the
young trees ascending, in large trees spreading wide, western part of the range also in winter; snow only
curved; branches of second order dense, spreading, at the highest elevations. It is mainly associated with
 Pinus strobiformis, P. pseudostrobus, P. ayacahuite. Picea crassifolia Kom., Bot. Mater. Gerb. Glavn.
Some other pines, Pseudotsuga menziesii var. glauca, Bot. Sada RSFSR 4: 177. 1923. Types: China, div. loc.,
Abies durangensis, A. vejarii (in Sierra Madre Ori- N. M. Przewalski s.n., Feb 1800; P. K. Kozlov s.n.,
ental), Cupressus lindleyi (= C. lusitanica Mill.), and May 1895 (syntypes LE).
C. arizonica may also occur. Broad-leaved trees are
e.g. Quercus castanea, Q. rugosa and Prunus serotina.
Etymology
The species epithet means ‘with thick leaves’.
Conservation
The (sub)populations of P. chihuahuana are widely
Vernacular names
566 scattered and very small, with a total of fewer than
100 to about 350 mature trees in each of the ca. 25 lo- Qinghai spruce; Qinghai yun shan (Chinese)
calities known. It is possible that other relict popula-
tions are hidden among the pine forests of the Sierra
Description
Madres of northern Mexico, yet its total area of oc-
cupancy (AOO) is unlikely to exceed 500 km2 and Trees to 20–25 m tall, d.b.h. to 50–60 cm; trunk mo-
is by all accounts much less than this; in addition it nopodial, straight; bark on trunk rough and scaly,
is severely fragmented. Unaware of its botanical sig- red brown. Branches of first order short, spread-
nificance, loggers have exploited this species where ing horizontally or upturned; branches of second
they have encountered it, reducing the number of order short, firm, spreading or ascending; crown
mature individuals. In many stands natural regen- pyramidal or (narrowly) conical, open in old trees.
eration has been observed to be poor or at best slow. Branchlets short, firm, leading shoots thick, pale
Awareness of its significance for conservation of bi- orange-yellow or greenish yellow, often pruinose,
odiversity is now increasing, but few subpopulations later grey, with prominent ridges and deep grooves,
are as yet under any effective protection. glabrous or with some scattered pubescence; pulvini
IUCN: EN (B1+2e) large, 2–2.5 mm, erect or recurved. Vegetative buds
ovoid-globose, more conical on leading shoots, up to
8–12 × 6–10 mm, not or slightly resinous; bud scales
Uses
triangular, acutish, keeled, with erose margins, or-
The rarity of this species renders it economically ange brown, often pruinose, persistent, leaving wide
unimportant as a timber tree. However, its poten- collars of perular scales at base of shoots. Leaves
tial size in old-growth stands has made it a valuable crowded above shoot, upper leaves directed forward
resource of good timber when encountered by log- and lower leaves curved upward, very rigid, (0.9–)
gers, and trees have been logged to be processed in 1.2–2.2(–2.5) cm long, 1.5–2.5(–3) mm wide, linear,
local sawmills. Spruces are uncommon to rare in curved or nearly straight, nearly quadrate-rhombic
northern Mexico, so there is, or was, an incentive in cross-section, keeled on two sides; apex obtuse-
to harvest them for their useful wood properties in acutish; amphistomatic, stomata in 4 bands; leaf col-
construction and carpentry. Introduction to the UK our bright green, with two whitish green stomatal
from a population in Nuevo Leon sampled in the bands. Pollen cones axillary, 1–1.5 cm long, yellowish
early 1980s appears to have been successful, but the pink. Seed cones terminal, erect at first, pendulous
species is still rare in cultivation. There are now a at maturity, sessile, ovoid-oblong or cylindrical, ob-
few more trees in several arboreta in the USA, the tuse at apex, (5–)7–11 cm long, 2.5–3.5 cm wide with
UK and Denmark from collections of seed made in opened scales; colour (immature) purplish red, ma-
Chihuahua in 1989 and the early 1990s (Grimshaw turing to green, with purple margins of seed scales,
& Bayton, 2009: 566). It is one of the most attractive ripe cones light brown or dark brown. Seed scales
species in this genus. broadly obovate-flabellate, slightly convex, flattened
when open, 1.5–2 × 1–1.7 cm at mid-cone; surface
smooth, usually finely striate, or slightly wrinkled,
glabrous; upper margin entire, slightly incurved,
rounded; base cuneate. Bracts rudimentary, ligulate,
2 mm, entirely included. Seeds ovoid-oblong, 3–3.5
mm long, brown; seed wings ovate-oblong, 10–13 ×
Description
4–5 mm, orange-brown.
Trees to 40–45(–50) m tall, d.b.h. to 1–1.5(–2) m;
trunk monopodial, straight; bark reddish brown,
Distribution
later grey with light brown and becoming rough
China: Gansu, Qinghai (Qilian Shan, around Qing- and fissured. Branches of first order short, slender,
hai Hu), Nei Mongol (Daqing Shan), Ningxia (He- spreading horizontally, curved upward at ends,
lan Shan). lower branches more pendant; branches of second
TDWG codes: 36 CHI-NX CHN-GS CHQ order short, dense, spreading or pendulous; crown
narrowly conical or narrowly columnar, especially
in the N or at high elevations, with branches to the 567
Ecology
ground or the bole free of branches for a third to
Picea crassifolia occurs in high mountain ranges half in dense forest stands. Branchlets slender, firm,
of Central Asia, mainly on N-facing slopes, above becoming pendulous, greenish yellow at first, soon
steppe or desert, at elevations between 1600 m yellowish brown, ridged and grooved, finely pu-
and 3800 m a.s.l. It occurs on calcareous and non-­ bescent when young; pulvini well developed, 2 mm
calcareous soils (the first soil type e.g. in Helan long, standing at nearly 90° to shoot axis. Vegeta-
Shan). The climate is cold continental and dry, with tive buds ovoid-conical, 5–6 mm long, resinous at
most of the precipitation falling as snow. It forms apex; bud scales obtuse-triangular, appressed, apices
mostly pure forests, here and there with Betula albo­ later spreading, red brown, persisting several years.
sinensis and groves of Populus tremula. Leaves spreading radially, crowded above leading
shoots, directed forward, (1.2–)1.5–2.5(–3) cm long,
(1–)1.5–2 mm wide, linear, straight or slightly curved,
Conservation
quadrate-rhombic in cross-section; apex acute (not
IUCN: LC pungent); amphistomatic, 2 narrow bands of 2–3
lines above, 2 bands of 4–6 lines below; leaf col-
our glaucous green. Pollen cones axillary, 1–1.5 cm
Uses
long, yellowish. Seed cones terminal, erect at first,
Qinghai spruce is probably only locally exploited for pendulous at maturity, sessile, ovoid-cylindric; apex
its timber (firewood?), as it occurs remote from any obtuse, (2.5–)3–6(–7.5) cm long, 2–2.5(–3.5) cm wide
major urban centres and even major roads. It is not with opened scales; colour (immature) green tinged
uncommon in cultivation in botanic gardens and with red, maturing to light reddish brown or pale
arboreta in Beijing, St. Petersburg and Moscow. In yellowish brown. Seed scales obovate-obtrullate,
the West it is now also appearing, but still very rare; thin and flexible, 1.2–1.5 × 0.9–1.2 cm at mid-cone;
some trees are possibly misidentified, e.g. as Picea abaxial surface smooth or finely striated, often un-
asperata. dulate, glabrous; upper margin rounded or truncate,
undulate, entire or erose-denticulate, sometimes
lacerate; base cuneate. Bracts broadly ovate, cuspi-
Picea engelmannii Parry ex Engelm., Trans. Acad. date, 3–6 mm long, entirely included. Seeds ovoid,
Sci. St. Louis 2: 212. 1863. 2–3 mm long, greyish brown; seed wings ovate-­
oblong, 10–12 × 4–5 mm, yellowish brown.
Etymology
Distribution
W. E. Parry named this species after the botanist
George Engelmann (1809–1884), but failed to pro- W North America: Rocky Mountains from British
vide a validating description, an omission made Columbia to North Mexico.
good by Engelmann. TDWG codes: 71 ABT BRC 73 COL IDA MNT ORE
WAS WYO 76 ARI CAL NEV UTA 77 NWM TEX 79
MXE-CU MXE-NL
Vernacular names
Engelmann spruce
 Ecology Description
Picea engelmannii is widespread in the Rocky Moun- Bark reddish brown, later grey with light brown.
tains, from 600 m to 3700(–4000) m a.s.l., the upper Leaves 1.5–2 mm wide. Bract scales of seed cones
limit progressively higher from N to S. It grows on 3–5 mm long.
various mountain soils, both calcareous and non-
calcareous. The climate is cold and humid (precipi-
Distribution
tation above 600 mm annually), with long, snowy
winters and short, cool summers. The species forms W North America: Rocky Mountains from British
extensive pure forests or mixed coniferous forests, Columbia to New Mexico.
568 with Abies lasiocarpa, Pinus spp., Pseudotsuga men­ TDWG codes: 71 ABT BRC 73 76 77
ziesii, Larix occidentalis, or Picea glauca as most
common associated species.
Conservation
IUCN: LC
Uses
Engelmann spruce is an important timber tree with Picea engelmannii Parry ex Engelm. subsp. mexi­
a high yield potential especially in managed stands cana (Martínez) P. A. Schmidt, Haussknechtia 4:
within its native range. Its knotty wood is not of very 38. 1988. Picea mexicana Martínez, Anales Inst.
high grade, but nevertheless increasingly used for Biol. Univ. Nac. México 32: 137. 1961; Picea engel­
home building, carpentry, furniture, plywood, and mannii Parry ex Engelm. var. mexicana (Martínez)
specialist uses such as musical instruments (pianos, Silba, Phytologia Mem. 7: 44. 1984. Type: Mexico:
violins). Uses for mining timber, railroad sleepers, Nuevo León, El Carmen, J. Vazquez 500a (holotype
and telephone poles have declined and mass pro- MEXU).
duction is now directed to the pulp wood industry,
especially in western Canada. Here massive clear cut
Description
operations still bare whole mountainsides regularly.
This species is rarely planted as an ornamental, al- This subspecies differs from the typical subspecies
though it will grow well even on poor soils, and only in its lighter (grey) bark, its narrower leaves (1–1.2
a few cultivars are known in the trade. Spruces are mm), and its narrower and slightly longer bract
not much in use as Christmas trees in North Amer- scales (4–6 mm).
ica, unlike in Europe where they are the commonest
genus for this purpose. The subspecies mexicana is
Taxonomic notes
rare in cultivation.
This taxon has first been found in the Sierra de la
2 subspecies are recognized: Marta, 75 km SE of Saltillo in NE Mexico, at 3000
m to 3400 m a.s.l. on moist, N-facing, steep slopes
Picea engelmannii Parry ex Engelm. subsp. engel­ of dolomite limestone. Taylor & Patterson (1980)
mannii. Picea glauca (Moench) Voss subsp. engel­ described a Picea “hybrida” population in S Chi-
mannii (Engelm.) T. M. C. Taylor, Madroño 15: 114. huahua (Cerro Mohinora). D. S. Correll & H. S. Gen­
1959; Picea glauca (Moench) Voss var. engelmannii try 23183 (MEXU) from that location (3100 m) has
(Engelm.) Boivin, Naturaliste Canad. 93: 272. 1966. cones (7 cm) resembling large cones of P. engel­
Type: USA: Wyoming, Wind River Mountains, mannii and leaves narrower than 1.3 mm, with an
F. V. Hayden s.n. (lectotype MO). acute (not pungent) apex. This material is best treat-
ed as P. engelmannii subsp. mexicana.
Picea engelmannii Parry ex Engelm. var. glabra
Goodman, Madroño 10: 177. 1950.
Distribution
Mexico: S Chihuahua, Nuevo León.
TDWG codes: 79 MXE-CU MXE-NL
 5–6 lines on lower side; leaf colour green or glaucous
Conservation
green above, two niveous white stomatal bands be-
The main population in the Sierra de la Marta (the low. Pollen cones axillary, 1.5–2.5 × 0.3–0.4 cm, yel-
type locality) was nearly exterminated in a forest fire lowish. Seed cones terminal, erect at first, pendulous
in 1975. Other populations, e.g. on Cerro Mohinora at maturity; peduncles oblique, 5 mm long; shape
in Chihuahua, are much smaller and declining, and ovoid-oblong to ellipsoid-cylindrical, obliquely ta-
even more susceptible to destructive fires. In ad- pering at base, obtuse at apex, (4–)6–12(–14) cm
dition, trees have been felled while regeneration is long, 2.5–4.5 cm wide, colour (immature) green
poor and slow. or purplish green, maturing to dark brown. Seed
IUCN: EN (A1a, B1+2c) scales obovate-oblong or obovate, slightly convex,
lateral margins recurved in opened cones, 1.2–2.2 569
× 0.8–1.5 cm at mid-cone; surface finely striated or
Picea farreri C. N. Page & Rushforth, Notes Roy. undulate and wrinkled, glabrous; upper margin ir-
Bot. Gard. Edinburgh 38: 130. 1980. Type: Myanmar regularly rounded, sometimes apically narrowed
[Burma]: Kachin, Myitkyina District, Feng Shui and recurved, entire; base cuneate. Bracts rudimen-
Ling (valley), R. Farrer 1435 (holotype E). tary, ligulate, 2–3 mm long, entirely included. Seeds
ovoid-oblong with narrowed apex, ca. 4 × 2.5 mm,
dark brown; seed wings ovate-oblong, 13–15 × 6 mm,
Etymology
lustrous orange-brown.
This species was named after Reginald Farrer, who
first collected it in the ‘Valley of the Winds and Wa-
Distribution
ter’ (Feng-Shui-Ling).
China, W Yunnan (Salween River); Myanmar [Bur-
ma] (Fen Shui Ling [valley & pass]).
Vernacular names
TDWG codes: 41 MYA
Farrer’s spruce
Ecology
Description
This species occurs in the mountains at elevations
Trees to 30–35 m tall, d.b.h. to 60–70 cm; trunk between 2400 m and 2700 m a.s.l., on limestone. The
mono­podial, straight or slightly curved; bark on climate is cool and wet, with heavy monsoon rains.
trunk rough and scaly, with small plates, grey or It forms small stands of pure spruce in usually open
greyish brown. Branches of first order long, slender, forest, with undergrowth of bamboo and juniper.
spreading horizontally or curved down, pendant A little higher occur Larix sp., Pinus cf. armandii
at the ends; branches of second order long, slen- and Tsuga dumosa, but at lower elevations broad-
der, pendulous; crown broadly conical. Branchlets leaved rainforest prevails, adding to the ecological
slender, drooping or pendulous, olive brown to pale isolation of Picea farreri (Page & Rushforth 1980).
orange-brown, with flat ridges and shallow grooves,
sparsely pubescent at first, soon glabrous; pulvini
Conservation
weakly developed, 0.6–1 mm long, at 40–60° from
shoot, pubescent, orange-brown. Vegetative buds This very rare and localized spruce was, until quite
ovoid, or ovoid-conical, on leading shoots 4–5 mm, recently, only known from a high mountain valley
on lateral shoots smaller, slightly resinous; bud in Myanmar [Burma] leading up to a pass cross-
scales obtuse, with erose margins, chestnut brown or ing into the deep valley of the Salween in Yunnan,
purplish brown, persisting several years. Leaves on China. It was then also found on that side of the
the upperside of shoots radially spreading forward, border, so now two or perhaps three (sub)popula-
more pectinate below, leaves subtending lateral buds tions are known to exist. Forest cutting and clearing
longest and at 90°, (1.5–)1.8–2.3(–2.5) cm long, 1–1.1 for agriculture are expanding into higher elevations,
mm wide, linear, straight or slightly curved, convex threatening the remaining montane coniferous for-
above, flattened and keeled below, acute and slightly est and therewith this rare species. The status of the
pungent at apex; epistomatic, stomata in 2 bands of Burmese population(s) remains poorly known as
 no foreign botanists have visited this remote valley ovoid-oblong, 5–7.5 cm long, 3–4 cm wide when
since the 1930s. opened; base flattened or tapered to a short pedun-
IUCN: EN (B1, 2a-c) cle; seed scales spreading widely, obovate, convex;
apex more or less rounded to slightly elongated, en-
tire or erose, usually incurved.
Uses
Farrer’s spruce may be used locally for construc-
Taxonomic notes
tion timber. Reginald Farrer introduced its seed to
England, where it was planted in several arboreta The spruces that occur in a broad zone between
and private parks. The resulting trees were believed Norway spruce (Picea abies) and Siberian spruce
570 to have been lost until a tree at Exbury Gardens in (P. obovata) have often been recognized as belong-
Hampshire, England was identified in 1979 by Chris ing to the latter species (provided that it was ac-
Page and Keith Rushforth as having grown from cepted as distinct at the species rank from P. abies).
seed under Farrer’s collection number 1435, the The extraordinary variability observed in the cone
same as the holotype specimen at the Herbarium of morphology of P. abies in Central and W Europe
the Royal Botanic Garden, Edinburgh (E). Herbari- (see e.g. Schmidt, 1991) appears to be absent in
um specimens from this tree were used to illustrate P. obovata in Siberia, as well as in the spruces of the
Picea farreri in my book Pinaceae (Farjon, 1990: 276) ‘transition zone’, the borders of which are poorly de-
and deposited in the Herbarium at the University of fined. However, certain character states, especially
Utrecht (U), now in the National Herbarium of the in the shape of the seed scales and their margins,
Netherlands. This tree subsequently died, but cut- are similar to P. abies, while the cones are mostly
tings were taken, rooted or grafted, and distributed smaller, as in P. obovata. The history of retreats of
among arboreta and pineta in England and abroad. the forests before the expansion of the Scandinavian
New plants were introduced in Scotland from ice cap during ice ages, and their subsequent return
sources in Yunnan, China recently, but this species in interglacials, can have caused an intermingling of
remains extremely rare in cultivation. these two closely related species in refugia, resulting
in a hybrid taxon, as is borne out by isozyme studies
(Schmidt, 2002a, b).
Picea ×fennica (Regel) Kom., Fl. SSSR 1: 145. 1934.
Pinus abies L. var. fennica Regel, Gartenfl. 12: 95, f.
Distribution
b. 1863; Picea obovata Ledeb. var. fennica (Regel)
A. Henry, in Elwes & Henry, Trees Gr. Brit. Ireland NE Russia: Archangelsk, Karelia, Kola Peninsula,
6: 1360. 1913. Type not designated. Komi; Finland?
TDWG codes: 14 RUN
Picea vulgaris Link var. uralensis Tepl., Bull. Soc.
Imp. Naturalistes Moscou 41: 249, f. 2, 3. 1869; Pi­
Ecology
cea ×fennica (Regel) Kom. subsp. uralensis (Tepl.)
P. A. Schmidt, Haussknechtia 4: 38. 1988. The dominant conifer in the northern taiga of Eu-
ropean Russia, in drier sites mixed or occasionally
replaced by Pinus sylvestris, with undergrowth of
Etymology
Vaccinium spp. or even lichens, in waterlogged sites
The nothospecies epithet refers to Finland (in a wid- by Sphagnum bogs. Occurring from near sea level
er sense than the present territory of that country). to 600–800 m a.s.l. depending on latitude, dwarfed
and becoming scattered at the Arctic tree line. Large
Vernacular names parts of its range are underlain by permafrost; the
deepest summer frost-free soils here are on river
Karelian spruce
floodplains, where Picea attains its largest size. Fre-
quently assosiated with Populus tremula and Salix
Description
spp., especially after fires and slowly replacing these
Trees; habit, bark and foliage similar to Picea abies pioneers in undisturbed sites, with natural cycles
of northern Scandinavia. Seed cones symmetrical, of forest fires ranging from 40 years to ­occasionally
400 years. On river floodplains Alnus glutinosa ovoid-globose, ca. 6 × 4–5 mm, not resinous; bud
characterizes a stadium between pioneer vegetation scales ovate, obtuse, recurved, light brown, shiny,
and returning spruce forest, especially in more sour- persisting several years. Leaves spreading radially
thern parts of the range of P. ×fennica. forward, dense, curved inward or nearly erect above
some shoots, parted below, 0.8–1.3(–2) cm long, 1.5
mm wide, linear, straight or curved, quadrangular,
Conservation
nearly equifacial; apex obtuse or acute; amphisto-
IUCN: LC matic, stomata in 4 narrow bands of 2–4 lines; leaf
colour glaucous green or bluish green. Pollen cones
axillary, numerous, 1–2 cm long, yellow. Seed cones
Uses
terminal, erect at first, pendulous at maturity, often 571
This spruce is an important timber tree; in the Komi very numerous in upper part of crown, short pe-
region of N Russia it is the mainstay of the econo- dunculate or sessile, ovoid-oblong or ovoid, often
my in a thinly populated region. The production of curved, (obliquely) tapering at base; apex narrowed,
timber amounted to 6.8 million m3 in the year 1940, obtuse, 3–6(–7) cm long, 1.5–2.5 cm wide with
a peak reached just before the Great Patriotic War opened scales; colour (immature) green or reddish
(WW II) commenced and diverted all manpower green, maturing to yellowish or orange-brown, old
to the fronts. Traditional houses there were all built cones light brown or red-brown. Seed scales obovate
with spruce logs and this use is making a come- or obovate-oblong, slightly convex, thin but not pa-
back in all of northern Russia. Much of the timber is pery, spreading at 40–60° from axis when opened,
processed to wood pulp for use in the paper indus- 1.2–1.5 × 0.8–1.2 cm at mid-cone; surface smooth,
try. Trees attributed to this hybrid species are in cul- finely striated towards apex, often with white resin
tivation mainly around the Baltic Sea and in Russia. dots, glabrous; upper margin rounded or truncate,
denticulate or entire; base cuneate. Bracts rudimen-
tary, ligulate, 2 mm, entirely included. Seeds ovoid,
Picea glauca (Moench) Voss, Mitt. Deutsch. 2–3 mm long, reddish brown or dark brown; seed
Dendrol. Ges. 1907 (16): 93. 1907. wings ovate-oblong, 6–8 × 4–5 mm, light orange-
brown.
Etymology
Distribution
The species epithet refers to the blue-green (glau-
cous) foliage. Boreal North America.
TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT PEI
Vernacular names
QUE 73 MNT 74 MIN SDA 75 MAI MIC NWH NWY
White spruce, Canadian spruce
Ecology
Description
This species is the major conifer in the vast boreal
Trees to 40–50 m tall, d.b.h. to 1–1.2 m; trunk mo- forest of North America; it occurs at elevations be-
nopodial, straight; bark soon flaking, yellowish tween 5 m and 1900 m a.s.l. (var. albertiana up to
green, turning grey brown, scaly and rough in large 2100 m). The soils are usually of fluvial or glacial
trees. Branches of first order short, slender, spreading origin, neutral or slightly acid and often podzolized.
or pendant; branches of second order short, dense; The climate is cold continental in much of its range,
crown (narrowly) conical or columnar, branches to but cold maritime in the extreme east, the precipi-
the ground, or bole free of (living) branches for one tation varies between 200 mm and 1250 mm, the
third in dense forest stands; Branchlets short, slen- growing season between 25 and 160 days. It grows
der, firm, whitish grey or buff, sometimes yellowish, in pure stands or mixed with (sparse) Betula papy­
finally grey; prominently ridged and grooved, gla- rifera, also with other conifers in various parts of the
brous or pubescent; pulvini small, sometimes with range; it invades successional stages with various
minute hairs, at 60–90° from shoot. Vegetative buds northern broad-leaved trees.
 TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
Uses
MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT PEI
Stretching accross the northern part of the North QUE 73 MNT 74 MIN SDA WIS 75 MAI MIC NWH
American continent White spruce is a major boreal NWY
timber resource. Its wood is strong and suitable for
general construction timber as well as pulp wood;
Conservation
the latter is probably its major use because in many
regions of its range sizes of trees are small in compar- IUCN: LC
ison to conifers growing further south. Log cabins in
Alaska are often built with White spruce, with the Picea glauca (Moench) Voss var. albertiana (S. Br.)
572 bark left on the outside and moss put between the Sarg., Bot. Gaz. (Crawfordsville) 67: 208. 1919.
logs. High grade, straight and even-grained timber Picea albertiana S. Br., Torreya 7: 126. 1907; Picea
is used for indoor finishing and flooring, carpentry glauca (Moench) Voss subsp. albertiana (S. Br.)
and joinery. Small volume uses are the manufacture P. A. Schmidt, Haussknechtia 4: 38. 1988. Type not
of boxes, musical instruments (sounding boards) designated.
and canoe paddles. White spruce is planted as an
amenity tree mainly in northern countries. In hor-
Description
ticulture, southern provenances (var. albertiana) are
more common and most of the dwarfed cultivars New shoots pubescent; leaves 1.5–2 cm long; apex
come from this source. Some dwarf cultivars origi- obtuse. Seed cones ovoid or ovoid-oblong, abruptly
nated from witches brooms and are particularly narrowing at base.
slow growing and suitable for rockeries.
Taxonomic notes
2 varieties are recognized:
This variety has been considered a product of intro-
Picea glauca (Moench) Voss var. glauca. Pinus gressive hybridization with Picea engelmannii.
glauca Moench, Verz. Ausländ. Bäume: 73. 1785.
Type not extant.
Distribution
Picea glauca (Moench) Voss var. densata L. H. Bai- Canada: SW Alberta; USA: Montana?
ley, Cult. Conif. N. Amer.: 108. 1933; Picea albertiana TDWG codes: 71 ABT
S. Br. var. densata (L. H. Bailey) W. L. Strong & Hills,
Canad. J. Bot. 84 (7): 1138. 2006.
Conservation
Picea glauca (Moench) Voss var. porsildii Raup, Sar-
gentia 6: 102, t. 12. 1947; Picea albertiana S. Br. var. IUCN: LC
porsildii (Raup) W. L. Strong & Hills, Canad. J. Bot.
84 (7): 1139. 2006.
Picea albertiana S. Br. subsp. ogilviei W. L. Strong & Picea glehnii (F. Schmidt) Mast., Gard. Chron.,
Hills, Canad. J. Bot. 84 (7): 1139. 2006. ser. 2, 13: 300. 1880. Abies glehnii F. Schmidt, Mém.
Acad. Imp. Sci. Saint-Pétersbourg, sér. 7, 12 (2):
176. 1868. Type: Russia: Russian Far East, Sakhalin,
Description
P. von Glehn s.n. (holotype LE).
New shoots glabrous; leaves 0.8–1.3 cm long; apex
acute, rarely obtuse. Seed cones ovoid-oblong, (ob-
Etymology
liquely) tapering at base.
This species commemorates Peter von Glehn (1835–
1876).
Distribution
Boreal North America: from Newfoundland and
Vernacular names
New York to NW Alaska and W Montana.
Sakhalin spruce; aka-matsu (Japanese)
 stricted area between 1000 m and 1100 m a.s.l. It
Description
usually grows on rocky, N- or NW-facing mountain
Trees to 30 m tall, d.b.h. to 60–70 cm; trunk mo- slopes with podzolic soils; it has also been recorded
nopodial, straight; bark soon flaking, reddish brown, from ultrabasic, heavy metal yielding volcanic soils.
becoming rough and scaly, breaking into irregular The climate is cold, moist maritime, with abundant
plates, grey-brown to purplish grey or dark grey. precipitation all year, increasing sharply with eleva-
Branches of first order long, slender, spreading tion to more than 1500 mm annually. It grows in pure
horizontally, but lower branches bent downward; stands, or mixed with Picea jezoensis and/or Abies
branches of second order dense, spreading horizon- sachalinensis; at lower elevations broad-leaved trees,
tally; crown pyramidal or conical. Branchlets short, e.g. Ulmus japonica, Tilia maximowicziana and Acer
slender, firm, orange or reddish brown, becoming pictum become important constituents of the forest 573
purplish brown; surface ridged and grooved, pu- in many places. Here Taxus cuspidata mixes with the
bescent in the grooves and on well developed, 1 mm Spruces and the broad-leaved trees.
long, densely set pulvini. Vegetative buds ovoid or
ovoid-conical, 4–6 × 4 mm, (slightly) resinous; bud
Conservation
scales acute-triangular, basal scales long cuspidate,
red-brown, shiny, persisting several years. Leaves IUCN: LC
radially spreading, the leaves above shoot directed
forward, more or less pectinate below, (0.6–)1–1.2
Uses
(–1.5) cm long, 1–1.5 mm wide, linear, curved or near-
ly straight, quadrate-rhombic or transversely broad- Sakhalin spruce is a timber tree of only local impor-
ly rhombic in cross-section; apex obtuse or (shortly) tance; its wood is used for construction and general
acute; stomata on all sides, in 1–2 lines above and carpentry. It has been introduced into cultivation
3–4 lines below on each face; leaf colour green or in Europe and North America, but remains largely
glaucous green, lower surfaces whitish green. Pol- confined to a few collections in botanic gardens and
len cones axillary, 1–2 cm long, yellowish. Seed cones arboreta, mostly in cool to cold winter regions.
terminal, erect at first, pendulous at maturity; ovoid-
oblong to cylindrical-oblong; apex often truncate
with opened scales, 3.5–8.5 cm long, 2.5–3.8 cm wide Picea jezoensis (Siebold & Zucc.) Carrière, Traité
with opened scales; colour (immature) purplish Gén. Conif.: 255. 1855.
green or dark violet, maturing to purplish brown
with greenish bands, old and ripe cones brown, basal
Etymology
part of scales dark purplish brown. Seed scales obo-
vate-oblong to broadly obovate, thin but rigid, 1.1–1.8 The species epithet refers to Yezo (Jezo), an alterna-
× 0.8–1.2 cm at mid-cone; surface striated, usually tive name for Hokkaido.
undulate, glabrous; upper margin undulate or emar-
ginate, erose-denticulate; base cuneate. Bracts rudi-
Vernacular names
mentary, ligulate, 2–3 mm long, entirely included.
Seeds ovoid-fusiform, 2.5–3 × 1.2–2 mm, light brown Yezo spruce; Ezo-matsu (Japanese); Jel ajanskaja
or yellowish brown; seed wings ovate-oblong, 7–10 × (Russian)
4–6 mm, yellowish brown or orange-brown.
Description
Distribution
Trees to 40–50 m tall, d.b.h. to 1–1.5 m; trunk mo-
Japan: Hokkaido, N Honshu; Russian Far East: S Sa-
nopodial, straight; bark on trunk rough and scaly,
khalin.
fissured, blackish brown or dark purplish grey.
TDWG codes: 31 SAK 38 JAP-HK JAP-HN
Branches of first order long, slender, spreading hori-
zontally; branches of second order slender, dense,
Ecology
drooping or pendant; crown pyramidal or broad
Picea glehnii occurs from near sea level in the north conical. Branchlets slender, firm, pale yellow at first,
to 1650 m a.s.l. on Hokkaido, on N Honshu in a re- later orange-yellow or reddish brown, shiny; ridged
 and grooved, glabrous or puberulent (form of Kam- ­ okkaido it occurs also in mixed coniferous-broad-
H
chatka); pulvini angular, 1 mm long, darker than 2nd leaved ­forests.
year shoot. Vegetative buds ovoid-conical, 5–8 ×
4–6 mm, not resinous; bud scales obtuse-triangular,
Uses
shiny orange-brown, persisting several years. Leaves
spreading radially, directed forward above shoot, Yezo spruce is an important timber tree in north-
pectinate below, 1–2(–2.4) cm long, 1.5–2(–2.2) mm ern Japan and the maritime provinces of NE China,
wide, linear, straight or slightly curved, shallowly Korea, and Russia. In several forest types it is a co-
obtriangular in cross-section, acute or mucronate at dominant over large areas where it is logged with
apex; epistomatic, stomata in 2 bands of 6–7 lines; other conifers in the forest such as larch and fir. Its
574 leaf colour shiny (dark) green above, 2 white sto- wood properties are similar to those of P. sitchen­
matal bands below. Pollen cones axillary, clustered sis, but this spruce does not attain the great dimen-
on pendulous shoots, 1.5–2 cm long, yellow. Seed sions of that species. Much of its wood is processed
cones terminal, erect at first, pendant at maturity, to pulp for the paper industry, but more specialized
often clustered, numerous, sessile or very short pe- uses are furniture making and (in Japan) musical in-
dunculate, cylindrical, sometimes slightly curved, struments. The Ainu string instrument tonkori has a
obtuse at apex, (3–)4–7(–9) cm long, 2–3.5 cm wide body made from Yezo Spruce. Log houses are con-
with opened scales; immature cones green, ripen- structed with its wood in northern parts of its range,
ing to light or darker yellowish or reddish brown, such as in Kamchatka where a small area of taiga is
sometimes pruinose. Seed scales numerous, small, dominated by this species. This spruce is commonly
opening wide, obovate-oblong to obtrullate or near- used in afforestation especially in Japan and Korea;
ly rhombic, very thin and papery, but rigid, 1–1.2 × in other parts of the Northern hemisphere it is less
0.6–0.8 cm at mid-cone; surface finely striated, un- often planted, either for forestry or for horticulture.
dulate or smooth and flat, glabrous; upper margin In western Europe, provenances from Honshu are
thin, undulate, erose-denticulate, flat or incurved; the only ones planted due to ‘late’ frost damage ex-
base cuneate. Bracts small, ligulate, with toothed perienced with trees from more northern sources.
and cuspidate apex, purplish, 4–5 mm long, in- Foresters have experimented with hybridization, e.g.
cluded. Seeds ovoid-cuneate, 3 × 2 mm, light brown; with P. glauca and P. mariana and with its closest
seed wings ovate-oblong, 6–10 × 4–5 mm, light or- relative, P. sitchensis. Only a few cultivars are known,
ange-brown or yellowish brown. usually sporting yellow new foliage.

2 subspecies and 2 varieties are recognized:

Distribution
NE China: coastal part of Jilin; Japan: Hokkaido, Picea jezoensis (Siebold & Zucc.) Carrière subsp.
Honshu; North Korea; Russian Far East. jezoensis var. jezoensis. Abies jezoensis Siebold &
TDWG codes: 31 KAM KHA KUR MAG PRM SAK 36 Zucc., Fl. Japon. 2 (2): 19, t. 110. 1842. Type: Japan:
CHM-JL 38 JAP-HK JAP-HN KOR-NK [“Abies No. 2, Coniferae ex insula Jezo”], P. F. von
Siebold s.n. (lectotype L).
Ecology
Picea ajanensis Fisch. ex Carrière, Traité Gén. Conif.:
Picea jezoensis occurs from near sea level to 2700 m 259–260. 1855; Picea jezoensis (Siebold & Zucc.) Car-
a.s.l. (subsp. hondoensis on Honshu: 1100 m to 2700 rière var. ajanensis (Fisch. ex Carrière) W. C. Cheng
m), on various (podzolic) soils. The climate is cold & L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 162. 1978.
temperate, moist or wet (1000 mm to 2500 mm an- Picea kamtchatkensis Lacass., Bull. Soc. Hist. Nat.
nual precipitation on Honshu). It is usually mixed Toulouse 58: 637. 1929; Picea jezoensis (Siebold &
with other conifers, e.g. Abies spp., Larix spp., Pi­ Zucc.) Carrière f. kamtchatkensis (Lacass.) S. L. Tung
nus spp. (with P. pumila in the north of the range), & Y. L. Chou, in Y. L. Chou, Lign. Fl. Heilongjiang:
and Tsuga diversifolia on Honshu, while Betula er­ 46. 1986.
manii is the most common broad-leaved tree. On Picea austromandshurica Silba, J. Int. Conifer Pre-
serv. Soc. 6 (2): 35. 1999.
 2 ­niveous white stomatal bands on the inverse-dor-
Description
sal side. Seed cones dark reddish brown; seed scales
New shoots pale yellow at first, later orange-yellow (broadly) obovate, nearly flat.
or yellowish brown; leaves 1.5–2 mm wide, only
slightly flattened (obtriangular in cross-section);
Distribution
stomatal band whitish. Seed cones 4–7(–9) cm long,
2–3.5 cm wide when opened; seed scales obovate- Japan: central Honshu.
oblong to obtrullate, very thin and papery but rigid; TDWG codes: 38 JAP-HN
surface undulate; margin erose-denticulate.
Conservation
575
Distribution
IUCN: LC
Japan: Hokkaido, N Honshu; Russian Far East.
TDWG codes: 31 KAM KHA KUR MAG PRM SAK 38
JAP-HK JAP-HN Picea koraiensis Nakai, Bot. Mag. (Tokyo) 33: 195.
1919.
Conservation
Etymology
IUCN: LC
This species was named after Korea, from where it
Picea jezoensis (Siebold & Zucc.) Carrière subsp. was first described.
jezoensis var. komarovii (V. N. Vassil.) W. C. Cheng
& L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 161. 1978. Picea
Vernacular names
komarovii V. N. Vassil., Bot. Zurn. (Moscow &
Leningrad) 35: 504. 1950. Type not designated. Korean spruce; Jel koreiskaya (Russian); Hongpi
yun shan (Chinese)
Description
Description
Seed cones 3–4 cm long, 2–2.2 cm wide when
opened; seed scales nearly rhombic, flat; upper mar- Trees to 30 m tall, d.b.h. to 60–80 cm; trunk mo-
gin strongly erose-denticulate. nopodial, straight; bark on trunk rough, scaly, fis-
sured below, dark grey-brown, newly exposed plates
brown. Branches of first order long, spreading hori-
Distribution
zontally, ends slightly assurgent; branches of second
China: Jilin (near coast); North Korea. order spreading, more pendant in older trees; crown
TDWG codes: 36 CHM-JL 38 KOR-NK pyramidal or conical. Branchlets slender, firm,
(pale) yellowish brown or pale red-brown, turning
grey brown, prominently ridged and grooved, gla-
Conservation
brous or rarely pubescent; pulvini small (1 mm), at
IUCN: LC 45–60° to shoot axis, darker. Vegetative buds ovoid
conical, 6–8 × 3.5–5 mm, resinous; bud scales trian-
Picea jezoensis (Siebold & Zucc.) Carrière subsp. gular-ovate, reddish brown, persisting several years.
hondoensis (Mayr) P. A. Schmidt, Haussknechtia Leaves spreading radially, directed slightly forward,
4: 38. 1988. Picea hondoensis Mayr, Monogr. Abiet. parted below shoot, 1.2–2.2(–2.5) cm long, 1.5–1.8 mm
Japan. Reich.: 51, t. 4, f. 9. 1890; Picea jezoensis wide, linear, straight or curved, transversely broadly
(Siebold & Zucc.) Carrière var. hondoensis (Mayr) rhombic in cross section; apex acute or obtuse; ca.
Rehd., Mitt. Deutsch. Dendrol. Ges. 1915 (24): 314. 4 lines of stomata on each face; leaf colour glaucous
1915. green or bluish green. Pollen cones axillary, crowded
near ends of shoots, 1.5–2.5 cm long, yellowish. Seed
Shoots reddish brown in the second year; leaves cones terminal, erect at first, pendulous at matu-
more or less flattened, 1.8–2.2 mm wide, with rity, often numerous, sessile, ovoid-oblong or short
 c­ ylindrical; apex obtuse, (4–)5–8 cm long, 2.5–4 cm some northern countries like Finland and Russia,
wide with opened scales; colour (immature) green but has not become a common forestry tree. It is in
or purplish green, ripening to light orange-brown or cultivation elsewhere, but uncommon and restricted
dull brown. Seed scales obovate-oblong to ellipsoid, to arboreta where some may be mis-identified as the
slightly convex, 1.3–1.9 × 1.1–1.6 cm at mid-cone; sur- similar P. koyamae from Japan.
face smooth, slightly striated, exposed part of abax-
ial surface shining, glabrous; upper margin entire or 2 varieties are recognized:
erose-denticulate, obtuse, rounded or truncate; base
cuneate, dark brown. Bracts rudimentary, ligulate, Picea koraiensis Nakai var. koraiensis. Types: North
2–4 mm, entirely included. Seeds ovoid-conical, 3–4 Korea: V. Komarov 82; T. Nakai 1880; T. Nakai s.n.
576 × 2–2.5 mm, dark brown; seed wings ovate-oblong, (syntypes LE, TI).
12–16 × 6–8 mm, pale yellowish, transparent.
Picea intercedens Nakai, J. Japan. Bot. 17: 4. 1941;
Picea koraiensis Nakai var. intercedens (Nakai)
Taxonomic notes
Y. L. Chou, Lign. Fl. Heilongjiang: 49. 1986.
This species is very similar to Picea koyamai and also
resembles P. obovata; with the latter introgressive
Description
hybridization is likely to occur. Schmidt-Vogt (1977)
recognized three varieties, described by Nakai (op. Seed cones short cylindrical, seed scales more or less
cit.) as species, of these only one is sufficiently dif- thick, obovate-oblong; upper margin entire, round-
ferent to be treated as a separate taxon. ed or obtuse.

Distribution Distribution
China: S. Heilongjiang, Jilin (Changbai Shan); North China: S Heilongjiang, Jilin (Changbai Shan); North
Korea; Russian Far East: Primorye (Ussuri River). Korea (Yalu River); Russian Far East: Primorye (Us-
TDWG codes: 31 PRM 36 CHM-HJ CHM-JL 38 KOR- suri River).
NK TDWG codes: 31 PRM 36 CHM-HJ CHM-JL 38 KOR-
NK
Ecology
Conservation
Picea koraiensis occurs in the mountains near the Sea
of Japan, at elevations between 1000 m and 1500 m IUCN: LC
a.s.l., on mountain slopes or (in the NE of its range)
along streams; on various (alluvial) soils. The climate Picea koraiensis Nakai var. pungsanensis (Uyeki
is cool, with snowy winters and annual precipitation ex Nakai) Farjon, Pinaceae (Regnum Veg. 121): 231.
above 1000 mm. It is usually mixed with other coni- 1990. Picea pungsanensis Uyeki ex Nakai, J. Japan.
fers, e.g. Abies nephrolepis, Pinus sibirica, and Larix Bot. 17: 3. 1941. Type: North Korea, Mt. Pung-san
gmelinii var. olgensis, merging with Picea obovata in [“Yotokurei”], H. Uyeki & T. Kondo s.n., 10 Sep 1935
the interior and to the north of its range. Along riv- (holotype MATSU? [“H. F. E. S.”]).
ers in the NE it is scattered in broad-leaved forest.
Description
Uses
Seed cones ovoid-oblong; seed scales thin, obovate-
Korean spruce is a valuable timber tree but its lim- ellipsoid or obtrullate; apex obtuse or truncate; up-
ited distribution compared to P. jezoensis makes it per margin erose-denticulate.
of minor importance economically. The wood is
mainly used for construction and for posts locally,
Distribution
but some of it may end up in the pulp mills together
with the much more common Yezo spruce. It has North Korea (Mt. Pung-san).
been introduced in arboreta and forestry trials in TDWG codes: 38 KOR-NK
 cm wide with opened scales; colour (immature)
Conservation
green or purplish green, ripening to dull brown or
IUCN: VU (D2) dark brown. Seed scales obovate to suborbicular,
slightly convex, often resinous, 0.6–1.8 × 0.6–1.2 cm
at mid-cone; abaxial surface smooth, finely striated,
Picea koyamae Shiras., Bot. Mag. (Tokyo) 27: 127. glabrous; upper margin rounded or truncate, den-
1913 [“koyamai”]. Type: Japan: Honshu, Nagano ticulate; base cuneate or slightly narrowed. Bracts
Pref., Yatsuga-dake, H. Koyama s.n. (syntype K). rudimentary, ligulate, 2–3 mm long, entirely in-
cluded. Seeds ovoid-conical, 2–4 × 1.5–2 mm, (dark)
brown; seed wings ovate-oblong, 5–10 × 3–5 mm,
Etymology
light yellowish brown. 577
This species has been named after the Japanese bot-
anist Mitsua Koyama, who discovered it in 1911 on
Distribution
Yatsuga-dake (mountain).
Japan: Honshu (Yatsugadake Mts.).
TDWG codes: 38 JAP-HN
Vernacular names
Koyama spruce; Yatsugadake-tohhi (Japanese)
Ecology
Picea koyamae occurs in small groves of 10 to 20
Description
trees (total population only a few hundred trees) on
Trees to 25 m tall, d.b.h. to 60 cm; trunk monopo- the N-facing slopes of mountains, at elevations be-
dial, straight; bark soon flaking, brown, turning tween 1500 m and 2000 m a.s.l. The rock is volcanic,
greyish purple, in large trees rough and scaly, dark the soil podzolized and slightly acid. The climate is
grey-brown or blackish grey. Branches of first or- cool, with cold, snowy winters and abundant pre-
der long, slender, spreading horizontally or assur- cipitation (1000 mm to 2000 mm annually). There
gent; branches of second order dense, covering the are frequent typhoons, which in the 20th century
main branches, finally pendant; crown pyramidal or have destroyed a substantial part of the population
conical, open and irregular in old trees. Branchlets (Schmidt-Vogt, 1977). There are some pure groves,
slender, firm, the longest flexible, (pale) yellowish but it is commonly mixed with Larix kaempferi, Pi­
brown, more orange-brown above, becoming darker cea alcoquiana var. acicularis, P. maximowiczii in a
with age; prominently ridged and deeply grooved, few localities, and various broad-leaved trees and
glabrous, or slightly pubescent in grooves on lateral shrubs.
shoots; pulvini well developed, 1–1.5 mm, on the up-
perside of the leading shoots erect. Vegetative buds
Conservation
ovoid-conical, covered by curved leaves, 4–13 × 3–7
mm, (very) resinous; bud scales triangular-ovate, This species is threatened with extinction under
light brown or red-brown, persisting several years, IUCN Red List criteria on the basis of its rarity
leaving a thick collar of perular scales at the base alone, regardless of whether the population has de-
of the shoot. Leaves spreading radially, on leading clined or is declining. It is considered that stochastic
shoots strongly assurgent or nearly erect, with lower events of destruction such as fire, storm, or landslide
leaves spreading and upper leaves hiding the shoot, could wipe out all existing trees in a short time if
rigid, (0.6–)0.8–1.5(–2) cm long, the longest leaves these happened in quick succession, compounding
near base of shoot, 1.5–2 mm wide, linear, curved, the damage beyond chance of recovery.
keeled on 4 sides, acutish at apex; with 3–5 lines of IUCN: EN (D)
stomata on each of 4 faces; leaf colour dark green
above, more glaucous green below. Pollen cones ax-
Uses
illary, clustered, 1–1.5 cm long, yellowish. Seed cones
terminal, erect at first, pendulous at maturity, ses- Due to its rarity and inaccessibility this spruce has
sile, ovoid-oblong or cylindric, obtuse or tapering no value as a timber tree; it grows slowly and is not
to a narrow apex (3–)4–9(–10?) cm long, 2.5–3.5(–4) useful in plantation forestry. In horticulture it has
 been successfully introduced in arboreta in the cool- rudimentary, ligulate, 2 mm long, entirely included.
er or colder NE of the USA and in northern Europe, Seeds ovoid-conical, 2–4 mm long, dark brown; seed
but it has not been grown commercially. wings ovate-oblong, 7–14 mm long, light brown.

Taxonomic notes
Picea likiangensis (Franch.) E. Pritz., Bot. Jahrb.
Syst. 29: 217. 1900. This is a highly variable species and several infraspe-
cific taxa have been recognized. Most of these were
at one time described as distinct species and there is
Etymology
apparently no strict consensus as to which entities to
578 The species epithet refers to the Lijiang Shan in Yun- recognize at what rank [e.g. Farjon, 1990: 287; Flora
nan, China, from where it was first described. of China 4: 29–30 (1999)].

Vernacular names Distribution

Likiang spruce; li jiang yun shan (Chinese) China: S Qinghai, S & W Sichuan, NW Yunnan,
SE Xizang [Tibet]; Bhutan.
TDWG codes: 36 CHC-SC CHC-YN CHQ CHT 40
Description
EHM-BH
Trees to 50 m tall, d.b.h. to 2–2.6 m; trunk mono­
podial, straight; bark rough and scaly, fissured, grey,
Ecology
with orange-brown freshly exposed bark. Branches
of first order long, slender, spreading or ascending; Both the type (type locality Lijiang Shan, Yunnan)
branches of second order variable, not pendulous; and the varieties are subalpine spruces (2900 m to
crown pyramidal or conical. Branchlets thick, rigid 4100 m a.s.l.) of the SW Plateau of China. They are
or slender, firm, pale yellowish grey or orange-brown usually associated with other conifers, e.g. Abies
to reddish brown; with prominent ridges and fine spp., Picea brachytyla, Larix potaninii, and Tsuga
grooves, pubescent or glabrous; pulvini 1–1.5 mm, spp. at the lower elevations.
at 60–90° to shoot axis. Vegetative buds ovoid-con-
ical or conical, 4–6 × 3–4 mm, (slightly) resinous;
Uses
bud scales small, triangular, appressed, red-brown
or purplish brown, persisting several years. Leaves Likiang spruce is a timber tree used for construc-
spreading radially, forward above shoot, parted be- tion, machines, poles, furniture, and wood pulp for
low, (0.6–)0.8–1.5(–1.7) cm long, 1–1.5(–2) mm wide, the paper industry. The bark is used to produce tan-
linear, straight or curved, rigid, quadrate-rhombic nin, resin is tapped or distilled from the wood, and
to transversely rhombic in cross-section; apex acute, the needles produce aromatic oils. In Europe and
pungent; amphistomatic, with fewer lines of stomata North America this species and its varieties can be
above than below; leaf colour dark green or glaucous found growing in arboreta and botanic gardens, as
green above, bluish green below. Pollen cones axil- well as in large private gardens with tree collections.
lary, 2–2.5 cm long, rose-red at first, yellowish at ma- The correct naming to variety of these trees is often
turity. Seed cones terminal, erect at first, pendulous at problematic.
maturity, often numerous, sessile or short, obliquely
pedunculate, oval-oblong, with oblique base and ob- 4 varieties are recognized:
tuse apex, (4–)7–12(–15) cm long, (3–)3.5–5 cm wide
with opened scales; colour (immature) magenta to Picea likiangensis (Franch.) E. Pritz. var. likiangen­
red, ripening to yellowish brown, reddish brown, sis. Abies likiangensis Franch., J. Bot. (Morot) 13 (8):
purplish brown or pale brown. Seed scales obovate 257. 1899. China: NW Yunnan, P. J. M. Delavay 1031
or broadly obtrullate, 1.5–2.6 × 1–1.7 cm at mid-cone; (holotype P). Fig. 189, 190, 191
abaxial surface smooth or striated, glabrous; upper
margin rounded obtuse, entire, or undulate, denticu- Picea likiangensis (Franch.) E. Pritz. var. bhutanica
late to repand-lacerate at apex; base cuneate. Bracts Silba, Phytologia 68: 41. 1990.
Picea likiangensis (Franch.) E. Pritz. var. forrestii
Conservation
Silba, Phytologia 68: 41. 1990.
The limited range of this variety is exacerbated by
forest destruction through logging and subsequent
Description
grazing of livestock, which has lead to a decline that
Shoots at first sparsely pubescent or glabrous; leaves is ongoing.
0.8–1.7 cm long, transversely rhombic in cross-sec- IUCN: VU (B1 b-c)
tion, 1–1.5 mm wide. Seed cones ovoid-oblong with
an oblique base, 7–12(–15) cm long, 3.5–5 cm whide Picea likiangensis (Franch.) E. Pritz. var. montigena
when opened; seed scales obovate or broadly obtrul- (Mast.) W. C. Cheng, in Chen, Taxon. Chin. Trees:
late; upper margin rounded-obtuse, entire, some- 40. 1937. Picea montigena Mast., Gard. Chron., 579
times denticulate and undulate. ser. 3, 39: 146. 1906. Type: China: Sichuan, Daxue
Shan, Kangding, [“China Occ. prope Tatien-lu”],
E. H. Wilson 3027 (holotype BM).
Distribution
China: W Sichuan, NW Yunnan, SE Xizang [Tibet];
Description
Bhutan.
TDWG codes: 36 CHC-SC CHC-YN CHT 40 EHM- Leaves very short (0.6–1.5 cm), quadrate rhombic.
BH
Distribution
Conservation
China: SW Sichuan (around Kangding, mainly to
IUCN: LC the W and S).
TDWG codes: 36 CHC-SC
Picea likiangensis (Franch.) E. Pritz. var. hirtella
(Rehd. & E. H. Wilson) W. C. Cheng, in Chen,
Ecology
Taxon. Chin. Trees: 40. 1937. Picea hirtella Rehd.
& E. H. Wilson, in Sargent, Pl. Wilson. 2: 32. 1914, This variety occurs in mountains above 3300 m a.s.l.
non Loudon (1838); Picea purpurea Mast. var. in coniferous forests.
hirtella (Rehd. & E. H. Wilson) Silba, Phytologia
68: 44. 1990. Type: China: Sichuan, Guan Xian,
Conservation
Banlan Shan, [“West of Kuan Hsien, Panlan Shan”],
E. H. Wilson 2084 (holotype A). This variety is believed to be of very limited extent of
occurrence (EOO) and under threat of deforestation
and logging.
Description
IUCN: EN (B1b)
Leaves slightly compressed, amphistomatic. Seed
cones yellowish brown at maturity; seed scales ob- Picea likiangensis (Franch.) E. Pritz. var. rubescens
trullate; margins denticulate. Rehd. & E. H. Wilson, in Sargent, Pl. Wilson. 2: 31.
1914. Type: China: Sichuan, Daxue Shan, Kangding,
[“Tachienlu & neighbourhood”], E. H. Wilson 2057
Distribution
(holotype A).
China: W Sichuan, SE Xizang [Tibet].
TDWG codes: 36 CHC-SC CHT Picea balfouriana Rehd. & E. H. Wilson, in Sargent,
Pl. Wilson. 2: 30. 1914; Picea likiangensis (Franch.)
E. Pritz. var. balfouriana (Rehd. & E. H. Wilson)
Ecology
Hillier, Rep. Conif. Conf. Roy. Hort. Soc. London
This variety occurs at high elevations between 3000 1931: 232. 1932; Picea purpurea Mast. var. ­balfouriana
m and 4000 m a.s.l. in coniferous forests. (Rehd. & E. H. Wilson) Silba, Phytologia 68: 44.
 1990; Picea likiangensis (Franch.) E. Pritz. subsp. shoot axis. Vegetative buds ovoid-conical, 5–9 × 4–5
balfouriana (Rehd. & E. H. Wilson) Rushforth, Int. mm, resinous only at base; bud scales small, trian-
Dendrol. Yearb. 1998: 61. 1999. gular, appressed but later free at apex, red-brown
or chestnut brown, persisting several years. Leaves
spreading radially, forward above shoot, widely
Description
parted below, 0.8–2(–2.5 or rarely 3 cm long at mid-
Shoots pubescent, orange or reddish brown; leaves dle part of shoot), 1–1.5 mm wide, linear, straight
1.5–2 mm wide. Seed cones 4–8 × 3–4 cm, often or curved, rigid, quadrate-rhombic to transversely
oblique, cleft at one side and purplish brown at rhombic in cross-section; apex hard pungent; episto-
maturity; seed scales thin, coriaceous, obtrullate, matic (rarely one or two intermittent lines on abax-
580 repand-lacerate. ial surface); leaf colour glossy green, greyish green
below. Pollen cones axillary, 2–2.5 cm long, rose-red
at first, yellowish at maturity. Seed cones terminal,
Distribution
erect at first, pendulous at maturity, often numerous,
China: S Qinghai, W Sichuan, SE Xizang [Tibet]. sessile or short, obliquely pedunculate, oval-oblong,
TDWG codes: 36 CHC-SC CHQ CHT with oblique base and obtuse apex, 5–12 cm long,
3–5 cm wide with opened scales; colour (immature)
purplish, ripening to brown or reddish brown. Seed
Conservation
scales thin, more or less flexible, obovate or rhom-
IUCN: LC bic, 1.5–2.2 × 1–1.5 cm at mid-cone; abaxial surface
smooth, glabrous; upper margin obtuse or constrict-
ed and incurved, often repand-lacerate at apex; base
Picea linzhiensis (W. C. Cheng & L. K. Fu) cuneate. Bracts rudimentary, ligulate, 2 mm long,
Rushforth, Int. Dendrol. Soc. Yearbook 2007: 48. entirely included. Seeds ovoid-conical, 2–4 mm
2008. Picea likiangensis (Franch.) E. Pritz. var. long, dark brown; seed wings ovate-oblong, 7–14
linzhiensis W. C. Cheng & L. K. Fu, Acta Phytotax. mm long, light brown.
Sin. 13 (4): 83. 1975. Type: China: Xizang [Tibet],
Zangbo River, Linzhi, G. X. Fu 676 (holotype PE).
Taxonomic notes
Recently, Rushforth (2008) reviewed the spruces
Etymology
occurring in the Zangbo [Yarlung Tsangpo] drain-
The species epithet refers to Linzhi (Linzhi Xian), age of SE Xizang [Tibet] and found that the spruces
China, the municipality within which the type spec- growing at high altitudes there differ consistently
imen was collected. from Picea likiangensis and its varieties known from
Sichuan and Yunnan, and have in fact some char-
acteres reminiscent of Picea spinulosa. This high
Vernacular names
altitude taxon was recognized as Picea likiangensis
lin zhi yun shan (Chinese) var. linzhiensis by Chinese botanists, and Rushforth
elevated it to species rank, which is here followed.
More collecting in areas outside this drainage in
Description
SW Sichuan and NW Yunnan, where it is reported
Trees to 50 m tall, d.b.h. to 2–2.2 m; trunk monopo- by the Chinese to occur, is needed to establish with
dial, straight; bark rough and scaly, silvery grey, with more certainty both its distribution and taxonomic
thin, brownish fissures. Branches of first order long, status.
slender, spreading or ascending; branches of second
order variable, not pendulous; crown columnar or
Distribution
conical. Branchlets thick, rigid or slender, firm, light
brown or slightly orange-brown, becoming greyish China: SW Sichuan, NW Yunnan (near Zhong­
in the second to fourth year, with prominent ridges dian?), SE Xizang [Tibet].
and fine grooves, brown pubescent with glandular TDWG codes: 36 CHC-SC CHC-YN CHT
hairs in the first year; pulvini 1 mm, at 60–90° to
 Ecology Description
In SE Xizang [Tibet] this species forms almost pure Trees to 21 m tall; trunk to 50 cm d.b.h. Leaves slight-
forests between 3000 m and 3800 m a.s.l., usually ly 4-angled (less so than in P. glauca), with whitish
well above a mixed coniferous forest belt in which stomatal lines on upper surface (similar to P. sitch­
Picea spinulosa is the dominant spruce. At around ensis). Seed cones (3–)4–7 cm long; seed scales short
3000 m it occurs also mixed with Pinus armandii, and more or less rounded, thin, with erose-denticu-
while at its upper limit it grows with Larix sp. and late upper margin, light brown.
Abies spp., the firs ultimately replacing the spruces
above 3600–3800 m a.s.l. Understorey trees include
Taxonomic notes
Betula szechuanica, B. utilis, Acer caudatum, Malus 581
baccata and Sorbus sp. and there may be a well de- A natural hybrid between Picea glauca and P. sitch­
veloped shrub layer with e.g. Rhododendron, Euony­ ensis, occurring where the parental species are sym-
mus, Lonicera, Borinda, and Enkianthus (Rushforth, patric.
2008).
Distribution
Conservation USA: Alaska (Kenai Peninsula).
TDWG codes: 70 ASK
IUCN: LC
Ecology
Uses
This taxon occurs in a transition between marine-
This species is a timber tree used for construction, coastal conifer forest dominated by Picea sitchensis
machines, poles, furniture and wood pulp for the and (upland) interior spruce forest dominated by
paper industry. The bark is used to produce tannin, P. glauca. Two of the commonest conifers to occur
resin is tapped or distilled from the wood and the with it are Tsuga heterophylla and T. mertensiana,
needles produce aromatic oils. In Europe and North the latter more in the interior of the Kenai Peninsula.
America this species could be found growing in ar-
boreta and botanic gardens from earlier, misidenti- Conservation
fied introductions. More recently, Keith Rushforth
IUCN: LC
has introduced this species in the UK from several
visits to Xizang [Tibet) between 1995 and 2000. The
Uses
identification given to trees of earlier introduction
that belong to this species, which were possibly in- This hybrid taxon is obviously being logged and
troduced by Francis Kingdon-Ward from the Zang- used as its parental species as and where it occurs.
bo (Tsangpo) Valley, is most likely Picea likiangensis. It has also attracted some attention from foresters
in plantation experimental gardens or plots outside
North America, e.g. in the UK and in SW Green-
Picea ×lutzii Little, J. Forest. (Washington) 51: 745. land, where several other northern species in the
1953. Type not designated. Pinaceae are being tested for plantation forestry.
Hybrids between species can grow more vigorously
than either of the parents.
Etymology
This nothospecies was named after H. J. Lutz of Yale
Picea mariana (Mill.) Britton et al., Prelim. Cat.
University, who discovered it in 1950–51.
Anth. Pter. New York: 71. 1888. Abies mariana
Mill., Gard. Dict., ed. 8: Abies No. 5. 1768. Type not
Vernacular names designated.
Lutz’s spruce
Etymology
The species epithet means ‘belonging to Marius’.
 Vernacular names Distribution
Black spruce, Bog spruce, Swamp spruce Boreal North America: from Newfoundland and
New Jersey to interior Alaska.
TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
Description
MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT PEI
Trees to 25–30 m tall, but in most of its range only QUE 74 MIN WIS 75 CNT MAI MAS MIC NWH PEN
5–10 m, d.b.h. to 50–60 cm; trunk monopodial, RHO VER
straight or curved; bark on trunk rough and scaly,
fissured, grey or blackish grey; inner bark brown.
Ecology
582 Branches of first order usually short, slender, pen-
dant; branches of second order short, dense, espe- Picea mariana occurs mostly in bogs or swamps and
cially in the top; crown variable, mostly narrowly on permafrost sites (‘muskeg’), at elevations between
conical or columnar, reaching to the ground, lower <150 m and 800 m a.s.l., occasionally in western
branches often layering. Branchlets slender, short, mountains to 1500 m or 1800 m a.s.l., on a variety of
yellowish, or reddish brown, especially in 2nd year, acid soils, often on peat, in the south predominantly
ridged and grooved, densely reddish brown pubes- so. The climate is cold subhumid, but with a wide
cent, glabrous in the 3rd year; pulvini small, dense- amplitude. Annual precipitation varies from 200 to
ly set in spirals, with light apices. Vegetative buds 1400 mm, the growing season from 25 to 160 days.
ovoid-conical; terminal buds 5–6 mm long, lateral Pure stands occur mostly on Sphagnum peat and
buds smaller, not resinous or slightly resinous; bud on permafrost, elsewhere it is usually mixed with
scales triangular, keeled and acute-cuspidate in ter- Picea glauca, Pinus banksiana and Abies balsamea;
minal buds, pubescent, purplish brown or purple, A. lasiocarpa and Pinus contorta in upland regions,
persisting several years. Leaves spreading radially, and Populus tremuloides after fire. In the SE of its
directed forward above shoot, parted below, (0.6–) range Picea mariana occurs in a mixed conifer-an-
0.8–1.2(–1.5) cm long, 0.7–0.8 mm wide, linear, but giosperm swamp type with Chamaecyparis thyoides,
slightly tapering towards an acute, pungent (but Larix laricina, Abies balsamea, Populus balsamifera,
sometimes nearly obtuse) apex, transversely rhom- Acer rubrum, Ulmus americana, Fraxinus nigra, and
bic in cross section; amphistomatic, with 1–2 lines of other broad-leaved species.
stomata on each face above, 3–4 lines below; leaf col-
our dark green to glaucous green above, green with
Conservation
bluish white stomatal bands below. Pollen cones ax-
illary, often very crowded, 1–1.5 cm long, yellowish IUCN: LC
brown. Seed cones terminal or subterminal, often
in great abundance, clustered in top of tree, short,
Uses
obliquely pedunculate or sessile, ovoid or (sub)
globular; base oblique or curved, (1.5–)2–3.5(–4) cm Black spruce is economically very important as a
long, 1.5–2(–2.8) cm wide with opened scales; colour source of pulpwood especially in the eastern parts of
(immature) reddish or (dark) violet, maturing to its great range. Its wood is light in weight and strong,
shiny red-brown or dark purple, old cones dark red- nearly white in colour, and contains relatively little
brown, grey-brown or blackish brown, usually per- resin. It is one of the few spruces in North America
sisting several years on the tree, finally deciduous. in use as a Christmas tree, due to its compact shape
Seed scales obovate-suborbicular, rigid, brittle at when taken from natural stands; this use is now in
last, 0.7–1.2 × 0.6–1 cm at mid-cone; surface usually decline. A ‘spruce beer’ beverage is brewed using
quite rough, shining, striated or wrinkled, glabrous; the needles, and young twigs and the needle resin
upper margin erose-denticulate, more or less undu- are also distilled for their aromatic properties to be
late, usually curved inward; base short, broad. Bracts used in cosmetics. There is anecdotal evidence that
rudimentary, ligulate, 1–2 mm, entirely included. the drinking of spruce beer saved the English inhab-
Seeds ovoid-cuneate, 2 mm long, blackish brown; itants of 18th century trading posts in Hudson Bay
seed wings ovate, 5–8 mm long, orange-brown. from succumbing to scurvy, caused by a deficiency
of vitamin C in sailor’s diets. Native Americans used Seed scales obovate, convex, opening very wide, 2–3
the split roots to bind together birch bark canoes, as × 1.8–2.5 cm at mid-cone; abaxial surface smooth,
their elastic properties tend to pull the seams tight. glabrous; upper margin denticulate; apex becoming
In horticulture Black spruce is valued for its com- reflexed; base cuneate. Bracts rudimentary, ligulate,
pact, slow growth and often glaucous leaves and a 4 mm long, entirely included. Seeds ovoid-oblong,
modest number of cultivars, both dwarf forms and 5–8 mm long, slightly flattened, brown; seed wings
variegated forms, are available in the trade. ovate-oblong, 16–23 × 6–9 mm, yellowish brown.

Taxonomic notes
Picea martinezii T. F. Patt., Sida 13 (2): 131. 1988.
Type: Mexico: Nuevo León, Montemorelos, 6 km Picea martinezii (i.e. the two pupulations in Nuevo 583
SE of La Trinidad, T. F. Patterson 5629 (holotype León) was at the time of their botanical discovery
MEXU). considered to be conspecific with P. chihuahuana,
to which the seed cones in particular are very simi-
lar (and unique among North American spruces).
Etymology
These disjunct populations were separated as a dis-
This species was named after Maximino Martínez tinct species based on some minor, but consistent
(1888–1964), who studied the conifers of Mexico. differences in the leaves and seed cone scales. Farjon
(1990: 241; 1998, [2001]) considered them conspecif-
ic with P. chihuahuana in the Sierra Madre Occiden-
Vernacular names
tal. However, recent analysis using DNA markers
Martinez’s spruce (Ledig et al., 2004) showed the two species to be less
closely related despite the morphological similari-
ties. Picea martinezii is therefore again accepted as
Description
a distinct species.
Trees to 30–35 m tall, d.b.h. to 0.8; trunk monopo-
dial, straight; bark on trunk rough and scaly, break-
Distribution
ing into numerous small plates, dark grey. Branches
of first order long, slender, in young trees ascending, Mexico: Nuevo Leon (Sierra Madre Oriental: Mon-
in mature trees spreading wide, curved; branches of temorelos, Zaragoza).
second order dense, spreading, becoming (partly) TDWG codes: 79 MXE-NL
pendulous; crown in large trees broad conical, open,
irregular. Branchlets firm, young shoots yellow, later
Ecology
reddish brown, finally grey, prominently ridged and
deeply grooved, glabrous; pulvini rounded, 1 mm, One locality with this species (near La Trinidad,
on leading shoots nearly erect. Vegetative buds Montemorelos) is described as on slopes of coarse
conical, with acute apex, 8–10 × 5–6 mm, slightly limestone talus in a sheltered canyon at 2100–2200
resinous; bud scales triangular-ovate, with scari- m a.s.l. in montane mixed coniferous-deciduous
ous margins, appressed, reddish brown, persisting broad-leaved forest. Picea martinezii is here asso-
3–4 years. Leaves spreading forward, parted below, ciated with Abies vejarii, Pinus spp., Taxus globosa,
1.6–2.7 cm long, 1–2 mm wide, linear, straight or and angiosperm broad-leaved trees such as Quercus
slightly incurved, flexible, slightly flattened, keeled spp., Tilia, Ostrya, Cornus, Ilex, Juglans, and Cratae­
on one or both surfaces, long acuminate-pungent at gus. There is no surface water, but frequent rain and
apex; amphistomatic, 4–10 lines of stomata on each fog provide ample moisture to sustain a lush meso-
side; leaf colour glossy green or greyish green. Pol- phytic forest type.
len cones near end of shoots, axillary, 1–1.5 cm long,
yellowish. Seed cones terminal, erect at first, pen-
Conservation
dulous at maturity, solitary or a few together, ses-
sile, cylindrical, with obtuse apex, 8.5–16 cm long, This extremely rare species is restricted to two pop-
4–6 cm wide with opened scales; colour (immature) ulations about 150 km apart. The largest population
bright green, ripening to shining (light) brown. has suffered some logging in recent years from ­local
 lumber companies; the smallest population has numerous, spreading or more pendant in old trees;
fewer than 15 mature trees. However, a considerable crown (broadly) conical, dense. Branchlets slender,
amount of regeneration has been observed in the flexible, yellowish brown or orange-brown, darker
largest population, despite reports of low seed vi- above, ridged and grooved, glabrous or puberu-
ability. A potential threat is forest fires, which could lent; pulvini small, numerous, sometimes darker
easily destroy the smallest population in one event. than shoot. Vegetative buds ovoid-conical or ovoid-
Awareness of the conservation issue is growing and globose, 2.5–4.5 mm long, not or slightly resinous;
local foresters are monitoring the situation, trying bud scales ovate or rounded, appressed, red-brown
to persuade loggers to leave these trees alone. In or purplish brown, persisting several years. Leaves
the 1980’s seed was collected and distributed to sev- spreading at 60–70° forward, those above shoot
584 eral arboreta and the Royal Botanic Gardens, Kew nearly appressed, parted below, on shaded shoots
(Rushforth, 1986). nearly pectinate, (0.6–)1–1.3(–1.6) cm long, 1–1.4
IUCN: CR (B1, 2c) mm wide, linear, straight or slightly curved, trans-
versely rhombic in cross-section, acutish to ob-
tuse at apex; amphistomatic, 2–4 lines of stomata
Uses
on each face; leaf colour lustrous dark green, with
This species has been logged for its timber by local whitish green stomatal lines. Pollen cones axillary,
sawmills and the wood has been used for construc- often numerous, 1–1.5 cm long, yellowish. Seed
tion purposes and carpentry. This use is now being cones terminal, erect at first, pendulous at matu-
discouraged for conservation reasons. It has been rity; peduncles short, oblique; shape ovoid-oblong
introduced to arboreta in Europe, Australia (Tasma- or oval-cylindrical; apex obtuse, (2.5–)4–6.5(–9?)
nia) and New Zealand; trees that still survive from cm long, 2.5–3.5 cm wide with opened scales; col-
this effort can be considered extremely rare. Some our (immature) green, ripening to light red-brown
may be misidentified as Picea chihuahuana, which or dull brown. Seed scales broadly obovate-cuneate,
has more glaucous, shorter leaves and entire, not near base of the cone suborbicular, 1.3–1.8 × 1–1.5 cm
denticulate seed scales. at mid-cone; surface striated or grooved longitudi-
nally, dull or shiny, often resinous, glabrous; upper
margin rounded or obtuse, entire or slightly erose;
Picea maximowiczii Regel ex Mast., Gard. Chron., base cuneate. Bracts rudimentary, ligulate, obtuse,
ser. 2, 13: 363. 1880. 2–3 mm long, entirely included. Seeds ovoid-­oblong,
(2.5–)3–4.5 × (1.5–)2.5–3 mm, dark brown or grey-
brown; seed wings ovate-oblong, 7–10 × 4–5 mm,
Etymology
yellowish brown or orange-brown.
This species was named after the Russian explorer
Carl Johann Maximowicz (1827–1891), who sent nu-
Distribution
merous seeds and plant specimens back to Europe
from Japan. Japan: Honshu, Nagano Prefecture (Fuji-yama, Fuji,
Yatsuga-dake).
TDWG codes: 38 JAP-HN
Vernacular names
Maximowicz’s spruce, Japanese bush spruce; Hime-
Ecology
baromomi (Japanese)
Picea maximowiczii is a rare species of the high
mountains in central Honshu, where it occurs at el-
Description
evations between 1100 m and 2000 m a.s.l. The soils
Trees to 20–25 m tall, often a low bush, d.b.h. to are derived from volcanic rock and usually podzolic.
50–60 cm; trunk monopodial, straight, or forked The climate is moist, with cool summers and cold
low above the ground; bark on trunk rough and winters, the annual precipitation ranges between
scaly, fissured below, grey-brown or grey. Branches 1000 mm and 2000 mm. It occurs in small, scattered
of first order long, slender, ascending or spreading groups, usually associated with Juniperus rigida,
more horizontally; branches of second order short, sometimes with Pinus densiflora and various broad-
leaved trees, mostly in very open, grassy terrain. The Picea maximowiczii Regel ex Mast. var. senanensis
variety senanensis has been found with Picea alco­ Hayashi, Ill. Useful Trees (Forest Trees): sub f. 43.
quiana and P. koyamae. 1969. Type not designated.

Uses Description
This small bushy tree has little value for timber and This variety differs from var. maximowiczii in its
is now protected from further exploitation. In Ja- shorter needles (0.6–1.3 cm) and smaller cones (2.5–
pan it is commonly planted in gardens, especially in 4.5 cm long), which have smaller, slightly pointed
Buddhist temple grounds, where it is valued for its (obtuse) seed scales and smaller seeds (2.5–3 × 1.5–2
dense habit and slow growth; these traits also make mm). 585
it a good but uncommon species for bonsai grow-
ing. Introductions to Europe and North America
Distribution
have mainly been of the var. senanensis, or perhaps
involved hybrids between the two varieties. In Euro- Japan: Honshu (Fuji, Yatsuga-dake).
pean horticulture it is not a specially valued spruce TDWG codes: 38 JAP-HN
and is mainly confined to arboreta and similar col-
lections of planted trees. No cultivars are known of
Conservation
this rare species.
IUCN: VU (B1+2c)
2 varieties are recognized:

Picea maximowiczii Regel ex Mast. var. maximo­ Picea meyeri Rehd. & E. H. Wilson, in Sargent,
wiczii. Type: Japan: Honshu, [locality not stated], Pl. Wilson. 2: 28. 1914. Type: China: Shanxi,
Tschonoski (Chônosuka Sugawa) s.n. (holotype K). Xiao Wutai Shan, Wutai Shan, F. N. Meyer 22672
(holotype A).
Description
Picea meyeri Rehd. & E. H. Wilson var. mongolica
Leaves (0.8–)1–1.3(–1.6) cm long. Seed cones (3.5–) H. Q. Wu, Bull. Bot. Res. North-East. Forest. Inst. 7
4–6.5(–9?) cm long; seed scales broadly obovate (2): 153. 1986.
with a cuneate base and a rounded, entire or slightly Picea meyeri Rehd. & E. H. Wilson var. pyramidalis
erose upper margin. Seeds 3–4.5 mm long, 2.5–3 mm H. W. Jen & C. G. Bai, J. Beijing Forest. Univ. 17 (1):
wide. 95. 1995; Picea meyeri Rehd. & E. H. Wilson f. py­
ramidalis (H. W. Jen & C. G. Bai) L. K. Fu & Nan Li,
Novon 7 (3): 262. 1997.
Distribution
Japan: Honshu (Fuji-yama).
Etymology
TDWG codes: 38 JAP-HN
This species commemorates the Dutch plant collec-
tor Frans N. Meijer (1875–1918), who travelled ex-
Conservation
tensively in Asia.
Despite its bushy habit (or is that a result?) this
spruce has been over-exploited and present popula-
Vernacular names
tions are scattered and small. It is a slow grower and
regeneration is often hampered by changes of land Meyer’s spruce; bai qian (Chinese)
use that have caused habitat degeneration. Most
trees are now situated in State Forests and would
Description
therefore enjoy some measure of protection.
IUCN: VU (B1+2c) Trees to 30 m tall, d.b.h. to 60 cm; trunk mono­
podial, straight; bark on trunk rough and scaly, with
papery flakes, grey. Branches of first order long,
 slender, spreading horizontally; branches of second earth, usually podzolized and non calcareous. The
order slender, numerous, thicker and shorter in top climate is cold, continental, especially in the west-
of tree; crown (broad) conical, or columnar in forest ern part of the range, with a moderate annual pre-
stands. Branchlets short, firm, colour variable, pale cipitation (500 to 800 mm). It grows in pure stands
yellow to light brown, with prominent ridges and or mixed with Picea wilsonii, Abies nephrolepis, and
deep grooves, young shoots variably pubescent or Larix gmelinii var. principis-rupprechtii, the latter at
glabrous, all glabrous in the 3rd year; pulvini well de- elevations above 2100 m a.s.l. on Wutai Shan.
veloped, 1.5–2 mm long, erect or slightly recurved,
darker than shoot. Vegetative buds ovoid-conical to
Conservation
broad conical, more or less acute, 6–10 × 4–8 mm,
586 resinous; bud scales triangular, acute, appressed, IUCN: LC
recurved near bud apex, light brown, persisting
several years. Leaves spreading radially, crowded
Uses
above shoot, directed forward, parted below, (0.8–)
1.3–2.5(–3) cm long, 2 mm wide, linear, curved, on Meyer’s spruce is an importan timber tree in north-
young trees straight, in cross-section transversely ern China, harvested both from natural stands and
rhombic, acutish at apex; amphistomatic, stomata from plantations. The wood is used for house build-
in 2–5 lines on each face above, in 4–5(–8) lines on ing and other construction, foot bridges, poles, fur-
each face below; leaf colour glaucous green or blu- niture making, and also, especially the plantation
ish green, stomatal bands bluish grey. Pollen cones timber, for wood pulp used in industrial manufac-
axillary, 2–2.5 cm long, reddish yellow, ripening to turing. This species is in cultivation for afforestation
orange-yellow. Seed cones terminal, erect at first, and as an ornamental tree in arboreta, parks, and
pendulous at maturity, (obliquely) short peduncu- large gardens, in China and (as a garden tree only)
late or sessile, oval-oblong to cylindrical, usually in Europe and North America.
very symmetrical, sometimes slightly curved, with
obtuse apex, (6–)7–10(–12) cm long, 2.5–3.5(–4) cm
wide with opened scales; colour (immature) pur- Picea morrisonicola Hayata, J. Coll. Sci. Imp.
plish red, maturing to (light) reddish brown or grey- Univ. Tokyo 25 (19): 220. 1908. Type: Taiwan:
brown. Seed scales obovate or suborbicular, 1.5–2 × Nantou, Chia-i Pref., Yu-Shan, [“Mt. Morrison”],
1–1.6 mm at mid-cone; surface finely striated, slight- T. Kawakami & U. Mori 2108 (lectotype TI). Fig.
ly convex, glabrous; upper margin entire, rounded, 192, 193
convex, in some cones denticulate or truncate; base
cuneate. Bracts ligulate-spathulate, 4–6 mm long,
Etymology
entirely included. Seeds ovoid-oblong, pointed at
base, 3–4 × 1.5–2.5 mm, brown or blackish brown; This species is named for Mt. Morrison (Yu-Shan),
seed wings obovate-oblong, 10–15 × 5–6.5 mm, yel- the highest mountain in Taiwan.
lowish brown or reddish brown.
Vernacular names
Distribution
Taiwan spruce, Mount Morrison spruce; Taiwan
China: S Gansu?, Hebei, Nei Monggol [Inner Mon- yun shan (Chinese)
golia], Shaanxi, Shanxi (Wutai Shan).
TDWG codes: 36 CHI-NM CHN-HB CHN-SA CHN-
Description
SX
Trees to 40–50 m tall, d.b.h. to 1.5 m; trunk mo-
nopodial, straight; bark on trunk rough and scaly,
Ecology
breaking into small, rounded plates with thin flakes,
Picea meyeri is a high montane to subalpine species, purplish grey or grey. Branches of first order long,
occurring at elevations between 1600 m and 2700 m massive, remote, spreading horizontally, curved
a.s.l., often restricted to the N-facing slopes of the down at the ends; branches of second order short,
mountains. The soils are partly mountain brown numerous and dense; crown broadly conical, in old
trees irregular and open. Branchlets slender, flexible, trees, e.g. Quercus variabilis, Acer insulare, and Bet­
the smallest new shoots only 2 mm thick, pale yel- ula sp. It does not appear to form large pure stands
low, underside nearly white, later grey, with small and is in fact often one of the less frequent conifer
ridges and narrow grooves, glabrous or rarely re- species in the mixed coniferous forests.
mote pubescent; pulvini small, reddish. Vegetative
buds ovoid-conical, 3–4 mm long, slightly resinous;
Conservation
bud scales triangular-ovate, obtuse, appressed, dark
red-brown, persisting several years. Leaves spread- Exploitation of indigenous old growth forests for
ing radially, pressed forward above shoot, parted timber has reduced the area of occupancy (AOO) of
below (leaves with axillary buds longest), perpen- this species considerably and the best stands of large
dicular, (0.8–)1–1.5(–2) cm long, (1–)1.5–2(–2.5) mm trees are now largely restricted to protected areas. 587
wide, linear, usually curved, longest leaves straight, IUCN: VU (A2c)
quadrate-rhombic or diamond shaped in cross-sec-
tion, acute or obtuse at apex; amphistomatic, 2 bands
Uses
of 2–3 lines above, 2 bands of 4–5 lines below; leaf
colour green or dark green, with fine, greenish white Taiwan spruce can attain large size and is a valuable
lines of stomata. Pollen cones axillary, 1–1.5 cm long, timber tree, but it has been over-exploited in the
yellow. Seed cones terminal, erect at first, pendulous past and is now relatively rare. The wood is used for
at maturity; peduncles short, oblique, or cones ses- construction, carpentry and furniture making. It was
sile; shape ovoid-oblong, tapered at base, obtuse at introduced to Europe and the USA by Ernest H. Wil-
apex, (4–)5–7 cm long, 2.5–3 cm wide with opened son in 1918, but has remained uncommon in cultiva-
scales; colour (immature) (greenish ) red or purplish tion, mostly found only in arboreta, botanic gardens
green, light brown or dull brown when ripe. Seed and large private tree collections, and trees still alive
scales obovate or suborbicular, opening wide, 1.2–1.5 today are often dating back to his first introduction.
× 1–1.2 cm at mid-cone; surface smooth or finely
striated, glabrous; upper margin entire, rounded or
truncate; base broad cuneate. Bracts rudimentary, Picea neoveitchii Mast., Gard. Chron., ser. 3, 33: 116,
ligulate, purplish, 2–3 mm long, entirely included. f. 50, 51. 1903. Type: China: W Hubei, E. H. Wilson
Seeds ovoid-oblong, 3–3.5 mm long, dark brown; 2601 (holotype BM). Pl. 24
seed wings ovate-oblong, 8–10 × 4–5 mm, light yel-
lowish or orange-brown.
Etymology
The species epithet commemorates James Veitch, a
Distribution
Chelsea nurseryman who sent E. H. Wilson out to
Taiwan (central mountains). China to collect plants; ‘neo-‘ refers to a new species
TDWG codes: 38 TAI named after Veitch.

Ecology Vernacular names

Picea morrisonicola is a high montane species, oc- Veitch’s spruce, Hubei spruce; da guo qing qian
curring at elevations between 2300 m and 3000 m (Chinese)
a.s.l. on N-facing mountain slopes and in ravines.
The soils are acid and podzolized. The climate is
Description
cold temperate, with a monsoon character and very
wet: annual precipitation exceeds 4000 mm. This Trees to 15–20 m tall, d.b.h. to 50–60 cm; trunk
spruce occurs in association with Tsuga chinen­ mono­podial, straight; bark on trunk rough and
sis, Pseudotsuga sinensis, Pinus armandii, and with scaly, exfoliating in small, thin flakes, grey. Branch-
Abies kawakamii and Juniperus squamata var. morri­ es of first order spreading, with upper branches
sonicola to the tree line; at lower elevations also with ascending, lower ones curved down; branches of
Chamaecyparis obtusa var. formosana, which can second order short, spreading or ascending; crown
form pure stands below 2500 m, and broad-leaved (broad) conical. Branchlets slender, firm, ­yellowish
 6

588

1
8

7 4

Pl ate 2 4. Picea neoveitchii. 1. Habit of tree. 2. Branchlet with leaves. 3, 4. Seed cones. 5. Seed scale with
seeds. 6. Seed scale. 7. Leaves. 8. Seeds.
brown or pale brown, turning grey, ridged and
Conservation
grooved, glabrous, or rarely sparsely pubescent; pul-
vini small, directed forward or nearly erect on lead- Extensive logging has reduced this species danger-
ing shoots. Vegetative buds conical or ovoid-conical, ously close to extinction in many areas; only a few
5–6 × 3.5–4 mm, not resinous; bud scales triangular, populations are within forest reserves. The Chinese
obtuse, brown, appressed, slightly pubescent at base. government has recently announced a policy that
Leaves spreading radially, curved and directed for- intends to halt large scale logging in western China,
ward above shoot, parted below, 1.5–2(–2.5) cm long, but the effects on the ground remain uncertain as in
ca. 2 mm wide, linear, usually curved, transversely many areas illegal or unregulated timber extraction
rhombic in cross-section, keeled; apex acute to pun- is common.
gent; amphistomatic, with 4–7 lines of stomata on IUCN: EN (B1a-b, 2a-b) 589
each face; leaf colour dark green, with faint, whitish
green stomatal lines. Pollen cones axillary, ca. 2 cm
Uses
long, yellowish. Seed cones terminal, erect at first,
pendulous at maturity, sessile, ovoid-oblong, mas- The wood of this timber tree is used for construc-
sive, with obtuse apex, (7–)8–14 cm long, 4.5–6.5 cm tion, poles, furniture, and wood pulp in the paper
wide with opened scales; colour (immature) green, industry. From E. H. Wilson’s herbarium specimens,
maturing to yellowish brown, ripe cones light brown. which include ripe cones, no trees seem to have
Seed scales few, large, very broadly obovate, or broad been grown, and its existence in cultivation outside
spathulate near cone apex, convex, with well defined China remains uncertain.
apophysis, opening wide, 2.5–3.2 × 2.5–3 cm at mid-
cone; abaxial surface finely striated, smooth when
ripe, with basal part more or less frayed; upper mar- Picea obovata Ledeb., Fl. Altaica 4: 201. 1833. Picea
gin entire, rounded or slightly obtuse, incurved, of- abies (L.) H. Karst. var. obovata (Ledeb.) Lindq.,
ten undulate. Bracts rudimentary, ligulate, 4–5 mm Acta Horti Berg. 14 (8): 307. 1948; Picea abies (L.)
long, entirely included. Seeds ovoid-oblong, 5–7 × H. Karst. subsp. obovata (Ledeb.) Hultén, Svensk
4.5 mm, dark brown; seed wings obovate, 15–18 × Bot. Tidskr. 43: 388. 1949. Type: Russia: Altai Mts.,
8–10 mm, light brown. C. F. von Ledebour et al. s.n. (holotype LE).

Distribution Etymology
China: Chongqing (Daba Shan), S Gansu, Henan The species epithet describes the shape of the seed
(Neixiang), NW Hubei, S Shaanxi, Shanxi (Wutai cone scales (obovate = inverse egg-shaped, i.e. with
Shan). the broadest pole at the apical end).
TDWG codes: 36 CHC-CQ CHC-HU CHN-GS CHN-
SA CHN-SX CHS-HE
Vernacular names
Siberian spruce; Jel sibirskaya (Russian)
Ecology
Picea neoveitchii occurs on medium-high moun-
Description
tains, at elevations between 1300 m and 2000 m
a.s.l., often on N-facing, steep, rocky slopes. The soils Trees to 40 m tall, d.b.h. to 1 m; trunk monopodi-
are non-calcareous, podzolic and acidic lithosols, al, straight; Branches of first order slender, long or
the species even grows in rocky talus. The climate short, spreading horizontally or curved downward;
is cold-temperate, more continental in the western bark grey at lower part of trunk, rough, scaly, break-
part of the range, with annual precipitation between ing into small plates. Branches of first order slen-
500–800 mm. In several areas only scattered trees der, long or short, spreading horizontally or curved
have been found, associated with Abies nephrolepis downward; branches of second order highly vari-
or Picea asperata; in Chongqing this species occurs able, spreading horizontally; crown pyramidal or
in coniferous forests with Abies fargesii, Picea asper­ narrowly conical. Branchlets slender, bright orange
ata, and P. purpurea. or red-brown, becoming grey, ridged and grooved,
 (sparsely) pubescent or glabrous; pulvini 1 mm, ob-
Ecology
lique. Vegetative buds ovoid-conical, 4–5 × 3–4 mm,
not or slightly resinous; bud scales triangular, ob- Picea obovata is a constituent of the boreal taiga of
tuse, light brown or reddish brown, persisting sev- northern Russia and Siberia, where it tends to domi-
eral years. Leaves spreading radially around shoot, nate on shallow soils over permafrost and occurs to
directed forward, parting below, 0.8–2.5 cm long, well within the Arctic Circle. In the southern parts
1–1.8 mm wide, linear, straight or curved, quadran- of its huge range it is forming almost pure forests or
gular in cross-section, acute at apex; amphistomatic, mixed with Abies sibirica in the Altai Mountains to
with 2–4 lines of stomata on each face; leaf colour ca. 2000 m a.s.l. In water-logged areas it becomes
(dark) green. Pollen cones in leaf axils, near end of a stunted, narrowly columnar tree and often grows
590 shoots, 1–1.5 cm long, yellow. Seed cones terminal, together with Larix gmelinii in the eastern part of its
at first erect, pendulous at maturity, sessile, cylin- range. Betula and Populus are common associated
drical, rarely ovoid-oblong, 4–8 cm long, 2.5–4 cm angiosperm trees in the more or less open conifer
wide with opened scales; colour (immature) green forests on slightly deeper and better drained soils. In
or red, maturing to brown or dark brown. Seed dryer soil situations Pinus sylvestris can grow with
scales obovate-oblong to flabellate, 1.5–2.5 × 1–2 cm the spruces, too. Picea obovata is extremely tolerant
at mid-cone; surface smooth, glabrous; upper mar- of low winter temperatures, withstanding extremes
gin obtuse or rounded, entire; base cuneate. Bracts below –60 °C with a totally frozen soil. Under such
rudimentary, ligulate, 2–3 mm, entirely included. extreme conditions, with short but often hot and dry
Seeds ovoid-oblong, 2–4 mm, dark brown or black- summers, it grows very slowly and trees with a trunk
ish brown; seed wings ovate-oblong or cuneate, (6– diam. of 10 cm can be a century or more old.
)10–15 mm long, light brown.
Conservation
Taxonomic notes
IUCN: LC
Picea obovata is considered a distinct species in Rus-
sia, but its range meets that of P. abies in NW Rus-
Uses
sia and certainly in Lapland on the Kola Peninsula
and in northern Finland. Here it is considered to be Siberian spruce is a major timber tree in Russia and
either a subspecies of P. abies, or the product of hy- represents the largest resource of standing timber by
bridization, Picea ×fennica (Regel) Kom., between volume in that country (and perhaps in the world).
the two taxa. That such hybridization has occurred Much of the wood is processed to pulp, but other
is more than likely with the repeated retreat of uses for which Picea wood is traditionally valued
P. abies into refuges that were situated in European also apply to P. obovata, including the careful con-
Russia during the ice ages. The two taxa mixed in struction of violins. In amenity planting it is less
their subsequent recolonisation of lost ground when prominent and only used commonly in parts of Rus-
the Scandinavian icecap retreated during warmer sia and rarely in other countries of eastern Europe.
interglacial phases. The seed cone morphology of
undisputed Siberian populations of P. obovata is
quite uniform and distinct from the larger but vari- Picea omorika (Pančić) Purk., Oesterr. Monatschr.
able cones of P. abies; it is where the two have histor- Forstwesen 27: 446. 1877. Pinus omorika Pančić,
ically been sympatric where identification becomes Neue Conif. Alp.: 4. 1876. [Gard. Chron., ser. 2, 7:
difficult. 620. 1877]. Type: Serbia: W Serbia, Rastiste, Crvena
Stena, J. Pančić s.n. (lectotype K, designated here).
Distribution
Etymology
Russia: from northern European Russia across Sibe-
ria to the Sea of Okhotsk, southward to Mongolia. Omorika is the local name, used as the species epi-
TDWG codes: 30 ALT BRY CTA IRK KRA TVA WSB thet by Pančić under Pinus.
YAK 31 AMU KHA MAG 37 MON
 TDWG codes: 13 YUG-SE
Vernacular names
Serbian spruce; Morika, Omorika (Serbian)
Ecology
Picea omorika occurs in the mountains of Serbia, at
Description
elevations between 300 m and 1700 m a.s.l., mainly
Trees to 30–40 m tall, d.b.h. to 0.8–1 m; trunk mono­ on N-facing slopes. The bedrock is Mesozoic lime-
podial, straight or curved at base; bark thin, rough and stone or rarely serpentine, the soils are mesic, humic
scaly, with papery flakes, (reddish) brown. Branch- and calcareous rendzinas, often very rocky. The cli-
es of first order numerous, short, slender, curved mate is cool montane, with moderately warm and
downward or pendant, assurgent at end; branches dry summers and snowy winters, with annual pre- 591
of second order slender, pendulous, hanging in cipitation around 1000 mm. The species is usually
dense sprays; crown narrowly conical or columnar, mixed with Picea abies, but Abies alba and Pinus
branches reaching to the ground. Branchlets short, ­nigra also occur, while Fagus orientalis and Castanea
slender, flexible, orange-brown or dull light brown, sativa are broad-leaved trees occurring at the lower
later grey-brown, with flat ridges and shallow, faint elevations, where the coniferous forest merges with
grooves, pubescent in 1st and 2nd year, then glabrous; the more extensive beech forest. It seems to be much
pulvini small, to 1.5 mm, nearly glabrous. Vegetative less competitive with Fagus than Abies alba or Picea
buds ovoid-conical, acute, lateral buds often numer- abies; however, it is capable of layering.
ous, 5–8 × 2.5–4.5 mm, not resinous or resinous at
base; bud scales ovate, acute, appressed, apices be-
Conservation
coming free, red-brown, turning orange-brown, per-
sisting several years. Leaves radially spreading at first, This species is a Tertiary relict known from fossil
but soon more flat and pectinate, with lower leaves evidence to have occurred in large parts of Europe
spreading at nearly 90° from shoot, (0.8–)1–2(–2.2) and to have been reduced to its present refugium on
cm long, 1.5–2.2 mm wide, linear, flattened, keeled the Balkan Peninsula by successive ice ages. Histori-
on both sides; apex obtuse or acutish; epistomatic, cally, its natural range has further decreased due to
stomata in two bands of 4–6 lines on lower surface; logging and fires, resulting in some 50 small relict
upper surface dark glossy green, sometimes glaucous stands, which together may not cover more than 60
green, two greenish white stomatal bands below. Pol- ha (Schmidt-Vogt, 1977). It is unclear to what ex-
len cones axillary, cylindrical, 2–2.5 cm long, yellow- tent the planting of Picea abies by foresters in the
ish. Seed cones terminal, often on small lateral shoots, region has contributed to the competition expe-
erect at first, pendulous at maturity; peduncles ob- rienced by P. omorika from this more agressively
lique or curved, scaly; cones ovoid-­oblong, oblique growing spruce. The two species do not seem to
at base; apex tapering, 4–6.5 cm long, 2–3 cm wide hybridize, but further planting of P. abies should be
with opened scales; colour (immature) dark purple, stopped and introduction of P. sitchensis, with which
maturing to purplish brown, ripe cones red-brown P. omorika does hybridize, must be prevented by all
or grey-brown. Seed scales suborbicular, thin, rigid, means.
convex, remaining imbricate for a long time, 1–1.5 IUCN: VU (D2)
× 1–1.3 cm at mid-cone; surface striated, sometimes
transversely undulate, often covered with dark resin,
Uses
glabrous; upper margin erose-denticulate. Bracts ru-
dimentary, ligulate, 2–3 mm long, entirely included. Serbian spruce is no longer logged for its timber.
Seeds ovoid, with acute apex, 3 × 2.5 mm, brown or On a limited scale it has been planted as a forestry
red-brown; seed wings ovate, 6–8 × 4–5 mm, light (timber) tree in Finland and Sweden. Its economic
red-brown or yellowish brown. value however is in horticulture, where it has been
widely used due to its characteristic narrow shape
with gracefully down-curving branches along the
Distribution
entire stem. It was first introduced in 1880–81 in
Balkan Peninsula: Serbia (Tara Mts., Drina River German and Swiss gardens, from where it rapidly
drainage). spread to other countries. It grows well on sandy
 soils and is tolerant to air pollution. It is mainly used below. Pollen cones axillary, 1–2 cm long, yellow-
as an ornamental tree, but in Denmark it has been ish. Seed cones terminal, erect at first, pendulous at
planted also as windbreaks. Serbian spruce will layer maturity, often very numerous, sessile, narrowly cy-
when planted in open situations and when there is lindrical, tapering towards apex, (4.5–)5–9(–10) cm
sufficient moisture. A number of cultivars has been long, 2–3.3 cm wide with opened scales; colour (im-
selected, especially dwarfed forms suitable for rock mature) green, purplish green or purple, ripening
and heather gardens, the latter en vogue in the 1960s to red-brown or dark (purplish) brown. Seed scales
and 1970s but later going out of fashion. broadly obovate or suborbicular, thin but rigid,
slightly convex, 1.2–1.7 × 1–1.4 cm at mid-cone; sur-
face finely striated, shining, undulate or flat, some-
592 Picea orientalis (L.) Peterm., Pflanzenreich: 235. times resinous, glabrous; upper margin rounded,
1838–45. Pinus orientalis L., Sp. Pl., ed. 2, 2: 1421. entire or lacerate; base cuneate. Bracts small, ligu-
1763. Type: Turkey: Trabzon, B. Balansa s.n. late, serrate at rounded apex, 5–6 mm long, entirely
(neotype K, designated by Farjon in Jarvis [2007: included. Seeds ovoid, pointed at apex, 3–4 × 2–2.5
745]). Fig. 194, 195 mm, red-brown or dark brown; seed wings ovate,
6–8 × 4–5 mm, orange-brown or yellowish brown.
Etymology
Distribution
The species epithet refers to its oriental origin (as
viewed from western Europe). Caucasus, N Turkey (coastal mountains).
TDWG codes: 33 NCS TCS 34 TUR
Vernacular names
Ecology
Oriental spruce, Caucasian spruce; Jel kavkasskaja,
Jel vostochnaya (Russian) Picea orientalis occurs in the mountains around the
eastern shore of the Black Sea, at elevations between
(400–)700–2100 m a.s.l., in Turkey mainly on the
Description
northern (seaward) slopes. It has a preference for
Trees to 40–50(–60) m tall, d.b.h. to 1.5–2.3 m; trunk acid soils. The climate is characterized by cool to
monopodial, straight; bark on trunk rough and cold winters (according to elevation) and relatively
scaly, breaking into small plates, brown or brown- warm, dry summers, the annual precipitation var-
grey. Branches of first order long, slender, spreading, ies between 1000 mm and 2000 mm; Picea occurs
curved downward, assurgent at end, but variable; generally on drier sites than Abies. It forms exten-
branches of second order long, slender, spreading sive pure stands, especially at higher elevations and
or pendant; crown pyramidal or conical to narrow- at the limit of trees, or is mixed with Abies nordman­
ly conical, dense, branches to the ground in most niana. The undergrowth is poor, Vaccinium being
trees. Branchlets short, slender, flexible, yellowish predominant. At lower elevations it occurs scattered
or pale brown, turning grey, with narrow ridges in broad-leaved forests, with Fagus orientalis, Acer
and grooves, short pubescent; pulvini small, apices spp., Ilex colchica and Taxus baccata.
lighter than shoot. Vegetative buds ovoid-conical,
acutish; terminal buds acute, 3–5 mm long, not res-
Conservation
inous; bud scales triangular, obtuse, appressed, but
apices more or less free, reddish brown, persisting IUCN: LC
several years. Leaves spreading radially, but pressed
against shoot above, parted and nearly pectinate be-
Uses
low, directed slightly forward, 0.6–0.8(–1) cm long,
0.7–1 mm wide, linear, straight, transversely rhom- Oriental spruce is an important timber tree in the
bic in cross-section, obtuse or obliquely acutish at Caucasus, where it forms extensive pure stands,
apex; amphistomatic, 1–2 lines of stomata on each many of which are managed for forestry. It has also
face above, 2–5 lines on each face below; leaf col- been introduced as a forestry plantation tree in
our dark glossy green above, whitish stomatal bands countries in the eastern Mediterranean. The wood
of this species is of good quality, comparable to that wide, linear, slightly curved, quadrangular in cross-
of Norway spruce, and is put to similar uses. Among section, with 4 prominent ribs, with acute to spines-
these are construction, flooring, carpentry, furni- cent (pungent), colourless apex; amphi­stomatic,
ture making, and parts of musical instruments. In 3–6 lines of stomata on each (grooved) face; leaf
horticulture, this spruce is sometimes grown as a colour green, glaucous green or bluish green. Pol-
Christmas tree, but more commonly as an amenity len cones axillary, cylindrical, 2–3 cm long, yellow.
tree for parks and large gardens in many European Seed cones terminal, erect at first, pendulous at ma-
countries and in the USA. A good number of culti- turity, numerous, concentrated in top of tree, sessile,
vars is in the trade, among which are dwarf forms, ovoid-oblong or cylindrical, obtuse or truncate at
forms with yellowish flushing leaves and those with apex, 5–8(–10) cm long, 3–4.5 cm wide with opened
‘mounding’ habits. The dense foliage with very small scales; colour (immature) green, ripening to yellow- 593
leaves, the bright red immature pollen cones, and ish brown or pale brown. Seed scales obtrullate or
the pendulous, purple (when still closed) seed cones broadly rhombic, very thin, papery but tough, undu-
make this an attractive species for large gardens. late, spreading wide, 1.8–2.4 × 1–1.5 cm at mid-cone;
surface smooth, finely striated, glabrous; upper mar-
gin very erose; apex undulate to emarginate; base
Picea pungens Engelm., Gard. Chron., ser. 2, 11: cuneate. Bracts rudimentary, ligulate-cuspidate, 3–5
334. 1879. Type: USA: Colorado, Clear Creek, mm long, entirely included. Seeds ovoid, pointed at
G. Engelmann s.n. (lectotype MO). apex, 3 × 2 mm, brown; seed wings obovate, 6–9 ×
5–6 mm, much shorter than the seed scale, yellow-
ish brown.
Etymology
The species epithet alludes to the sharp pointed
Distribution
(pungent) needles.
USA: Rocky Mountains, mainly in SE Idaho, Wyo-
ming, Utah and Colorado, with isolated populations
Vernacular names
in Montana, Arizona and New Mexico.
Blue spruce, Colorado spruce TDWG codes: 73 COL IDA MNT WYO 76 ARI UTA
77 NWM
Description
Ecology
Trees to 40–50 m tall, d.b.h. to 1–1.5 m; trunk mo-
nopodial, straight; bark rough and scaly, deeply Picea pungens is a subalpine species occurring in the
fissured at lower part of trunk, dark grey-brown. Rocky Mountains at elevations between 1800 m and
Branches of first order numerous, moderately long 3300 m a.s.l., commonly along mountain streams or
or short, spreading horizontally, more erect or as- on moist, N-facing slopes. The soils are mountain
surgent near top; branches of second order dense, lithosols and streambed gravels of various origin,
rigid, spreading horizontally; crown conical, in old usually poorly developed. The climate is continental,
trees broad columnar below the top, branches com- with long, cold and snowy winters (frost free days
monly to the ground. Branchlets slender, firm, yel- 60 or less) and short, but relatively warm summers.
lowish brown to orange-brown, darkest above, later Annual precipitation ranges from 600 mm to 900
turning grey, prominently ridged and grooved, gla- mm. This species grows in small, scattered groves,
brous; pulvini small, directed forward, slightly dark- especially near perennial streams, or scattered and
er than shoot. Vegetative buds ovoid-oblong, obtuse, mixed with Pinus contorta, Pseudotsuga menziesii
5–8 × 3–4 mm, not resinous; bud scales triangular, var. glauca, or Populus tremuloides. It is everywhere
obtuse, loosely appressed, apices recurved, yellow- a rare constituent of the subalpine forest.
ish brown or pale brown, basal scales keeled, acute,
persistent, forming collars at base of shoots. Leaves
Conservation
spreading radially, rigid, directed upward and for-
ward, parted below shoot, 1.5–3 cm long, 1–1.5 mm IUCN: LC
 4 × 3 mm, resinous; bud scales triangular, obtuse,
Uses
appressed, shiny dark chestnut brown, persisting
Blue spruce is widely planted as an ornamental; several years. Leaves above shoot closely appressed,
perhaps the most famous trees of this species are covering it entirely, with lower leaves more or less
planted in front of Lenin’s mausoleum in the Krem- parting, all directed forward, with leaves on shaded
lin in Moscow. Its symmetrical crown and glaucous shoots more remote and spreading, 0.7–1.4 cm long,
blue foliage make it an ideal tree for gardening and 1.5–1.8 mm wide, linear, straight or curved at base
landscaping. In nature trees with green foliage exist, only, more or less dorsiventrally flattened, keeled on
but these are not much wanted as the stock for horti- both sides, obtuse-mucronate at apex; usually epi­
culture; instead blue forms have been repeatedly se- stomatic, with two bands of densely set stomata be-
594 lected. A substantial number of cultivars, both tree low, but sometimes 1–2 intermittent lines of stomata
forms and dwarf forms, have been raised displaying above; leaf colour bright green or glossy dark green
various colours and hues of the needles which tend above, two greenish white stomatal bands below.
to change with their age and become less intense. It Pollen cones axillary, conical, 1.5–2.5 cm long, light
is also popular as a Christmas tree, especially in the red, ripening to rose or yellowish. Seed cones ter-
USA. The wood of this species is brittle and knotty minal, erect at first, pendulous at maturity, sessile,
because it retains its branches on the bole and is ovoid to ovoid-oblong, tapering to base, obtuse at
therefore of little commercial value. apex, 2.5–5(–7) cm long, 1.7–3 cm wide with opened
scales; colour violet, purple or bright crimson when
immature, ripening to purplish brown or dull (light)
Picea purpurea Mast., J. Linn. Soc., Bot. 37: 418. brown. Seed scales rhombic, papery at apex, curved,
1906. Picea likiangensis (Franch.) E. Pritz. var. undulate, spreading wide when ripe, 1–1.5 × 0.8–1.2
purpurea (Mast.) Dallim. & A. B. Jacks., Handb. cm at mid-cone; abaxial surface smooth or finely
Conif.: 334. 1923. Type: China: Sichuan, Min striated, often resinous, glabrous; upper margin
River, Songpan, [“ad Sung Pan prope Tibetam”], thin, elongated, undulate, incurved in some cones,
E. H. Wilson 3026 (holotype BM). erose-denticulate; base shortly narrowed. Bracts
rudimentary, ligulate, 1–2 mm long, entirely includ-
ed. Seeds ovate-oblong, pointed at base, 2.5–3 mm
Etymology
long, (dark) purplish brown; seed wings (obliquely)
The species epithet refers to the purple colour of the ovate-oblong, 5–7 × 3–4 mm, orange-yellowish.
seed cones.
Taxonomic notes
Vernacular names
This species is quite unlike P. likiangensis (Franch.)
Purple-cone spruce; ziguo yun shan (Chinese) Pritzel, to which it has been linked as a variety in sev-
eral works describing (Chinese) conifers (e.g. Dal-
limore & Jackson, 1966; Wright, 1955; Den Ouden
Description
& Boom, 1965). It is undoubtedly closely allied to
Trees to 40–50 m tall, d.b.h. to 1–2 m; trunk mo- P. ­likiangensis and its varieties and has been placed
nopodial, straight; bark on trunk rough and scaly, with that aggregate group in the subsection Sitch­
grey-brown, with freshly exposed parts of bark or- enses E. Murray. The leaves of the Chinese mem-
ange. Branches of first order spreading horizontally, bers of this subsection of spruces vary from nearly
with lower branches curved down; branches of sec- equi­facial amphistomatic (P. likiangensis var. liki­
ond order short, slender, dense, spreading or more angensis) to nearly invers-dorsiventral epi­stomatic
pendant; crown pyramidal or narrowly conical, trees (P. purpurea).
at high elevations making narrow spires. Branchlets
slender, flexible, very numerous, pale (pinkish) yel-
Distribution
low-grey, finely grooved and ridged, young shoots
densely pubescent; pulvini small, directed forward China: Gansu, E Qinghai, NW Sichuan.
at 30–50° from shoot. Vegetative buds conical, ca. TDWG codes: 36 CHC-SC CHN-GS CHQ
 Picea retroflexa Mast., J. Linn. Soc., Bot. 37: 420.
Ecology
1906. Picea asperata Mast. var. retroflexa (Mast.)
Picea purpurea is a subalpine species of continental W. C. Cheng, in Chen, Taxon. Chin. Trees: 38.
mountains, occurring in a spruce belt at elevations 1937; Picea aurantiaca Mast. var. retroflexa (Mast.)
between 2600 m and 3600 m a.s.l., predominantly C. T. Kuan & L. J. Zhou, Fl. Sichuan. 2: 71. 1983.
on N-facing slopes. The soils are either grey-brown Type: China: Sichuan, Daxue Shan, near Kangding,
mountain soils or lithosols, usually podzolic. The [“Tachien-lu”], E. H. Wilson 3030 (holotype A).
climate is cold continental, with low to moderate
precipitation, most of it as winter snow. It grows in
Etymology
pure forests or mixed with several other species of
Pinaceae, e.g. Picea asperata, P. wilsonii, P. neo­veitchii The species epithet (Latin retroflexus = ‘bent back, 595
(rare), Larix potaninii, and Abies fargesii, which pre- reflexed’) refers to the pulvini on the shoots, which
vails above the spruce belt towards the tree line, and Masters described as being “patenti-reflexi”.
with some broad-leaved trees, usually Populus spp.
and Betula spp. in clearings. At lower elevations
Vernacular names
Tsuga chinensis and Quercus spp. may occur.
Tapao Shan spruce; linpi yun shan (Chinese)
Conservation
Description
IUCN: LC
Trees to 40–45 m tall, d.b.h. to 1–1.5 m; trunk mono­
podial, straight; bark on trunk rough and scaly,
Uses
breaking into small, flaking plates, grey or brownish
This species yields high quality timber used for grey; inner bark yellowish. Branches of first order
construction, furniture making, poles, machine short, numerous, spreading horizontally; branches
and instrument making, including musical instru- of second order short, rigid, numerous, spreading
ments, and to a limited extent for pulp in indus- laterally; crown narrowly conical or columnar, espe-
trial manufacturing, e.g. paper. It was introduced to cially in trees at high altitude. Branchlets short, firm,
England early in the 20th century by Ernest Wilson thick, light brown or orange-brown, prominently
and Joseph Rock and is commonly found growing ridged and deeply grooved, glabrous or often ferru-
in arboreta both in Europe and North America, but ginous pubescent; pulvini strongly developed, 1–2 ×
sometimes misidentified as P. likiangensis, or treated 1–1.5 mm, on strong shoots almost erect. Vegetative
as a variety of it (for the UK presumably based on buds broadly conical, closely surrounded by curved
its treatment in Dallimore & Jackson’s Handbook, leaves, 5–10 × 5–10 mm, resinous, often pubescent
1966). According to Rushforth (1987) introductions at base; bud scales triangular, keeled, appressed, or-
by Wilson from W Sichuan grow to taller, more ange-brown or with purplish apex, persisting several
columnar trees than those from Rock’s collections, years, leaving broad collars of perular scales at shoot
originating from S Gansu, where the climate is drier bases. Leaves spreading radially, curved forward,
and colder in winter. (1–)1.2–1.8(–2.5) cm long, (1.2–)1.5–2 mm wide, lin-
ear, curved, quadrangular or transversely rhombic
in cross-section, with prominent ribs; apex pungent;
amphistomatic, on upper surface 2 bands of 2–3
lines, on lower surface 2 bands of 4–6 lines of sto-
mata; leaf colour light green or glaucous green. Pol-
len cones axillary, 2 3–5 cm long, reddish, ripening
to reddish yellow. Seed cones terminal, erect at first,
then pendulous, sessile, oval-oblong to cylindric-
conical; apex obtuse, 8–13 cm long, 2.5–4 cm wide
with opened scales; immature cones purplish red,
maturing to purplish or reddish brown, ­ripening
 to lustrous brown. Seed scales obovate-oblong or
Uses
slightly obtrullate, those near base suborbicular,
spreading wide when ripe, 1.5–2 × 1.2–1.5 cm at mid- Although no uses are specifically reported of this
cone; abaxial surface striated, shining, more or less species, its timber has been exploited together with
convex, glabrous; upper margin rounded or obtuse, that of other species in the area and put to the same
slightly erose-denticulate, incurved, straight or uses. This spruce was introduced to Europe and the
slightly reflexed when opened base cuneate. Bracts USA by Ernest Wilson and is still present in several
ligulate-lanceolate, 5–6 mm long, entirely included. arboreta, often identified as P. asperata. Accord-
Seeds ovoid-oblong, 3–4 mm long, dark brown or ing to Rushforth (1987), trees in cultivation (in the
red-brown; seed wings obovate-oblong, 10–15 × 5–7 UK) are “noticeably greener than Dragon spruce”
596 mm, pale brown or yellowish brown. (P. asperata), which could indicate that Wilson in-
troduced seed from a green-leaved form of this
species. As Wilson himself noted (on specimen),
Taxonomic notes
glaucous forms also occur in its native habitat. This
In Flora of China 4: 28 (1999) this species has been ‘mixture’ of green and glaucous forms is common
made a synonym of P. asperata var. asperata, while among conifers in all families and has given rise to
other (Chinese) works (e.g. Ying et al., 2004; Far- both valuable horticultural varieties and much taxo-
jon, 1990, 1998, [2001]) have maintained the species nomic confusion.
rank, or included it as a variety of P. aurantiaca (Flo-
ra of Sichuan). A re-examination of relevant collec-
tions and populations seems desirable; this should Picea rubens Sarg., Silva N. Amer. 12: 33. 1899.
include work on DNA sequences. Type: Illustration in A. B. Lambert, Descr. Pinus
vol. 1, t. 28 (as Pinus rubra). 1803. (lectotype).
Distribution
Etymology
China: W Sichuan.
TDWG codes: 36 CHC-SC The species epithet means ‘red’ and describes the
colour of the cones (when young?).
Ecology
Vernacular names
Picea retroflexa is a typical subalpine species, which
occurs between 3000 m and 4000 m a.s.l. (to 4700 m Red spruce, Eastern spruce; Epinette rouge (French)
E of Dawu, Schmidt-Vogt, 1977), mainly on N-fac-
ing slopes on acidic soils. The climate is continental
Description
alpine with low annual precipitation. At the highest
elevations it grows either pure or mixed with Abies Trees to (25–)30–35(–50) m tall, d.b.h. to 1–1.5 m;
squamata, but at lower elevations Picea likiangensis trunk monopodial, straight; bark soon rough and
var. rubescens, P. aurantiaca, Abies chensiensis and flaking, reddish brown or purplish grey with brown,
Tsuga chinensis may occur with it. Betula albosinen­ in old trees fissured near base of trunk. Branches
sis is the only common broad-leaved tree species in of first order long, slender, spreading horizontally,
these forests. sometimes slightly assurgent; branches of second
order long, slender, spreading laterally; crown nar-
rowly pyramidal or conical. Branchlets slender,
Conservation
flexible, light yellowish brown at first, later turn-
The limited range of this taxon, in conjunction with ing orange or reddish brown, ridged and grooved,
exploitation of timber trees in the subalpine conifer the young shoots more or less densely pubescent,
forests in which it occurs, have led the Conifer Spe- but soon glabrous; pulvini small, pubescent at base.
cialist Group to infer past and possibly ongoing de- Vegetative buds ovoid-conical, acute, 5–8 mm long,
cline. Due to difficulties with identification it would slightly resinous; bud scales triangular, the outer
be hard to quantify this decline accurately. scales acute and reflexed, red-brown or chestnut
IUCN: VU (B1c) brown, persisting several years. Leaves spreading
r­ adially but parted below, directed obliquely for- leaved trees can be common or dominant, especially
ward, on strong shoots more or less assurgent above on better soils.
shoot, 1–1.5(–1.7) cm long, ca. 1 mm wide, narrow-
ly linear, usually slightly curved, quadrangular in
Conservation
cross-section, acute or nearly obtuse at apex; am-
phistomatic, stomata in 2–4 lines on all four faces; IUCN: LC
leaf colour shiny light green. Pollen cones axillary,
1.5–2.5 cm long, reddish at first, yellowish when ripe.
Uses
Seed cones terminal, erect at first, pendulous at ma-
turity, short pedunculate or sessile, ovoid, tapering Red spruce is an important timber tree of medium
towards base, obtuse at apex, 2.5–5(–6) cm long, 1.8– size but with a relatively limited natural range. Its 597
3.5 cm wide with opened scales; colour (immature) wood is light in weight and cream coloured, strong
purplish red or green with a purple tinge, maturing and straight grained. It has varied applications and,
to red-brown or purplish brown, old cones lustrous besides the mass use for paper pulp, more special-
red-brown. Seed scales obovate-cuneate, convex, ized uses include construction, boat building, flag
0.8–1.5 × 0.9–1.3 cm at mid-cone; surface slightly poles, cooperage, and especially musical instru-
striated or nearly smooth, often with white resin ments. Spruce wood is ideal for sounding boards
dots, glabrous; upper margin entire (in old cones and bodies of string instruments, from pianos (the
erose), rounded or obtuse, incurved; base cuneate. keys hit strings) to guitars and violins. It is not of-
Bracts rudimentary, ligulate, ca. 2 mm long, entirely ten possible to ascertain which species of spruce has
included. Seeds ovoid-oblong, 2–3 × 1.5 mm, light been used, as anatomically the wood of three species
brown or dark brown; seed wings obovate, 5–9 × 3–5 growing in this area is indistinguishable. Red spruce
mm, light brown or orange-brown. is uncommon in cultivation probably because it is
not very distinct and few cultivars are known. Lo-
cally the resin from trunk wounds has been used as
Distribution
chewing gum, perhaps learned by early settlers in
E Canada: maritime provinces, extreme SE Ontario, Maine from native tribes. This use has been redun-
S Quebec; USA: New England States and Appala- dant for nearly a century as other sources of gum
chian Mountains. replaced it.
TDWG codes: 72 NBR NFL-SP NSC ONT PEI QUE
75 CNT MAI MAS NWH NWJ NWY PEN VER WVA 78
NCA TEN VRG Picea schrenkiana Fisch. & C. A. Mey., Bull. Sci.
Acad. Imp. Sci. Saint-Pétersbourg 10: 254. 1842.
Ecology
Etymology
Picea rubens occurs from near sea level on the coasts
of the maritime provinces of Canada, to the higher This species was named after Alexander Gustav von
slopes of the Appalachian Mountains (1100 m to 1500 Schrenk (1816–1876), a German botanist working in
m a.s.l.). In the NE lowland areas the species grows Russia.
mainly on acid soils (pH 4–5.5) of alluvial origin, in
the mountains also on acidic, peaty or rocky soils
Vernacular names
generally unfavorable for most of the other tree spe-
cies of NE North America. It is climatically restrict- Schrenk’s spruce, Asian spruce; Jel Schrenka (Rus-
ed to areas with a cool, moist oceanic climate, with sian); Xuelingshan (Chinese)
annual precipitation between 875 mm and 2000 mm
(increasing with elevation). It is commonly mixed
Description
with Picea glauca or Abies balsamea, more rarely
with Picea mariana, which occupies swamps and Trees to 40–50(–60) m tall, d.b.h. to 1–2 m; trunk
bogs but may extend to drier sites. Rare or local as- monopodial, straight; bark on trunk scaly, with
sociated conifers are Abies fraseri, Tsuga canadensis, small plates, blackish grey; inner bark orange.
Pinus strobus, and Chamaecyparis thyoides. Broad- Branches of first order short, numerous, spreading
 and descending; branches of second order short, nu- is cold continental. It forms usually pure forests, but
merous, very dense, usually spreading; crown nar- it is sometimes mixed with Abies sibirica (A. sibirica
rowly conical or columnar, dense, branches to the subsp. semenovii with Picea schrenkiana subsp. tian­
ground in most trees. Branchlets short, thick, rigid, schanica), at lower elevations with Ulmus and Popu­
pale yellowish or yellowish grey, prominently ridged lus along streams. Juniperus pseudosabina occurs
and deeply grooved, variously pubescent or gla- usually on S-facing slopes outside the spruce forest,
brous; pulvini prominent, assurgent, 1.5–2 mm long. but may also form a shrub cover in it.
Vegetative buds conical, broad, sometimes ovoid,
acute, with crowded pulvini at base, 5–10 × 4–7 mm,
Uses
not resinous; bud scales triangular, appressed, shiny
598 yellowish brown, persisting several years. Leaves Schrenk’s spruce produces valuable timber in large
spreading radially, directed forward, (1–)2–3(–3.5) volumes and quantities, but its exploitation has
cm long, 1–1.5(–2) mm wide, linear, curved, trans- been hampered by the remoteness of the mountains
versely rhombic in cross-section, with 2 ribs on op- where it occurs, far from ports and industrial cen-
posite sides; apex pungent; amphistomatic, with 2–4 tres. It is a magnificent tree, but it is quite rare in cul-
lines of stomata in each of 4 grooved faces; leaf col- tivation outside Russia and even there not used very
our green, with whitish stomatal lines. Pollen cones often. A few trees are planted in arboreta in Europe
axillary, 1.5–2.5 cm long, yellowish red, yellow when and the USA; this species should also be used more
ripe. Seed cones terminal, erect at first, pendulous often in parks and large gardens. A few compact
at maturity, shortly pedunculate or sessile, cylin- ‘dwarf ’ forms are known as cultivars, mostly grown
drical-oblong, sometimes slightly tapering towards in Central and E Europe.
the obtuse-truncate apex, (6–)8–10(–12) cm long,
2.5–3.5(–4) cm wide with opened scales; colour (im- 2 subspecies are recognized:
mature) purplish or greenish, ripening to purplish
black or dull brown. Seed scales obovate, broad, Picea schrenkiana Fisch. & C. A. Mey. subsp.
opening at 90°, 1.3–1.8 × 1.2–1.5 cm at mid-cone; schrenkiana. Type: Kazakhstan: Dzhungarskiy
abaxial surface striated or wrinkled, glabrous; upper Alatau, Songaria, Cljekirgo Pass, A. G. von Schrenk
margin rounded or truncate, slightly incurved, en- s.n. (lectotype LE).
tire or erose. Bracts rudimentary, ligulate, 2–3 mm
long, entirely included. Seeds ovoid, pointed at base, Picea prostrata Isakov, Fl. Kirgiz. SSR 10: 374. 1962.
4–5 × 3 mm, dark brown with whitish spots; seed
wings ovate, 8–10 × 5–6 mm, orange-brown.
Description
Leaves (1.5–)2–3(–3.5) cm long, 1–1.5 mm wide. Seed
Distribution
cones cylindrical-oblong; seed scales at mid-cone
China: Xinjiang (Tian Shan); Kazakhstan, Kir- 12–15 mm wide; abaxial surface striated or nearly
gyzstan (mountains around Naryn River). smooth.
TDWG codes: 32 KAZ KGZ 36 CHX
Distribution
Ecology
Kazakhstan.
A high montane to subalpine species of Central TDWG codes: 32 KAZ
Asia, occurring between 1300 m and 3000(–3600)
m a.s.l., especially on N-facing slopes and in cool
Conservation
ravines. It grows on various mountain soils, usually
in rocky places with seepage water from snowmelt IUCN: LC
(perpetual snow at higher elevations). The climate
Picea schrenkiana Fisch. & C. A. Mey. subsp. tian­ base; bark on trunk scaly, breaking into rough plates,
schanica (Rupr.) Bykov, Izv. Akad. Nauk Kazahsk. dark grey; inner bark brown. Branches of first or-
SSR., ser. Bot. 5: 22. 1950. Picea tianschanica Rupr., der long, spreading horizontally; branches of second
Mém. Acad. Imp. Sci. Saint-Pétersbourg, sér. 7, order long, slender, spreading or pendant; crown
14 (4): 72. 1869; Picea morinda Link subsp. tian­ pyramidal or broad conical, in old, large trees with
schanica (Rupr.) Berezin, Bot. Zurn. (Moscow & abundant reiteration. Branchlets slender, flexible,
Leningrad) 50: 493. 1970; Picea schrenkiana Fisch. pale brown to almost white on the underside, turn-
& C. A. Mey. var. tianschanica (Rupr.) W. C. Cheng ing yellowish or orange-brown, shallowly ridged and
& S. H. Fu, Fl. Reipubl. Pop. Sin. 7: 146. 1978. Type grooved, shiny, glabrous; pulvini 1–1.5 mm, at nearly
not designated. Fig. 196, 197 90° from shoot. Vegetative buds ovoid-conical, acute
or obtuse, 4–5 × 2–3 mm, slightly resinous at base or 599
not resinous; bud scales triangular, obtuse, appressed,
Description
light brown, persisting several years and becoming
Leaves (1–)1.5–2(–2.5) cm long, 1.4–2 mm wide. Seed dark brown. Leaves spreading radially, stiff, on high-
cones elliptical-oblong, with broad, wrinkled seed er branches pressed forward above shoot, pectinate
scales. Schmidt-Vogt (1977) who treated this taxon below the shoot, 1.5–2(–2.5) cm long, ca. 1 mm wide,
as a species, reported cones 10–20 cm long (com- narrowly linear, usually curved, almost quadrangu-
monly 12 cm), but specimens seen from LE and PE lar in cross-section, keeled, pungent at apex; stomata
had cones only 9–10 cm long and cones of similar mainly on two (invers dorsal) faces below, none or
size were seen during a field trip to Kirgyzstan by a few faint lines above; leaf colour dark or bright
the author in August 2000. green, with two silvery white stomatal bands. Pol-
len cones axillary, crowded, 2–3.5 cm long, rose-red
at first, yellowish at maturity. Seed cones terminal,
Distribution
erect at first, later pendulous; peduncles short, ob-
China: Xinjiang (Tian Shan); Kirgyzstan (moun- lique or curved; shape of cones cylindrical-oblong,
tains around Naryn River) obtuse at apex, (4.5–)5–9(–10) cm long, (1.5–)2–3
TDWG codes: 32 KGZ 36 CHX (–4) cm wide with opened scales; colour (immature)
yellowish green or green, ripening to pale brown or
yellowish brown. Seed scales rhombic to obtrullate,
Conservation
sometimes irregular, very thin, papery, 1–1.5 × 0.6–1
IUCN: LC cm at mid-cone; surface finely striated, undulate,
sometimes with dark spots, glabrous; margins often
recurved; apex irregularly dentate or lacerate. Bracts
Picea sitchensis (Bong.) Carrière, Traité Gén. Con- small, ligulate lanceolate, 5–8 mm long (sometimes
if.: 260. 1855 [“sitkaensis”]. Pinus sitchensis Bong., nearly half as long as seed scale), included, but often
Mém. Acad. Imp. Sci. Saint-Petersbourg, sér. 6, Sci. visible with wide opened seed scales. Seeds ovoid,
Math. 2: 164. 1832. Type not designated. Fig. 198 2–3.5 × 1.5–3 mm, light or dark brown; seed wings
ovate-oblong, 6–10 × 4–5 mm, light yellowish.
Etymology
Distribution
The species epithet means ‘from Sitka’ a small port
in Alaska, USA. Pacific Coast Region of North America: from Alaska
to California.
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
Vernacular names
Sitka spruce
Ecology
Picea sitchensis occurs from tidewater up to steep
Description
mountain sides in Alaska and British Columbia,
Trees to 60–85(–90) m tall, d.b.h. to 4–4.5(–5) m; generally to ca. 900 m a.s.l. (highest record 1189 m),
trunk monopodial, straight, old trees buttressed at always in proximity to oceanic weather. The soils
 are variable, usually with a thick humus layer. The
Etymology
climate is very humid, annual precipitation ranges
from 1300 mm to 3750 mm, the winters are moder- This species was named after James Edward Smith
ate (in coastal Alaska snow in winter usually stays (1759–1828), founder and first President of the Lin-
only above 200 m). On Vancouver Island and on the nean Society of London.
Olympic Peninsula in Washington this spruce attains
its greatest size. It is usually mixed with Tsuga hetero­
Vernacular names
phylla (shade tolerant competitor), Pseudotsuga men­
ziesii and Thuja plicata, other associated conifers are Indian spruce, West Himalayan spruce, Morinda
Chamaecyparis lawsoniana (locally), Xanthocyparis spruce; rai, rewari, salla, ragha, morinda (Hima-
600 nootkatensis, and Abies amabilis, at higher elevations laya); changye yunshan (Chinese)
replaced by Tsuga mertensiana or A. lasiocarpa; Al­
nus rubra alongside rivers and Acer macrophyllum in
Description
groves are common broad-leaved trees.
Trees to 50(–60) m tall, d.b.h. to 1.5–2.5 m; trunk
monopodial, straight; bark becoming rough, scaly,
Conservation
greyish brown to grey, breaking into irregular plates.
IUCN: LC Branches of first order long, slender, spreading hori-
zontally, often assurgent at ends; branches of sec-
ond order long, slender, dense, strongly pendulous;
Uses
crown conical, old trees broad columnar. Branchlets
Sitka spruce grows to the largest tree of its genus and long, slender, flexible, pale yellowish brown or grey-
is abundant in the coastal forests between roughly ish brown, prominently ridged and grooved, gla-
43° and 62° N along the Pacific Ocean. It is a highly brous; pulvini well developed, 1.5 mm long, directed
valuable timber tree with growth rates exceeding forward at 45° from shoot, brown. Vegetative buds
those of other species and, in old growth stands, ovoid conical, (5–)8–12 mm long, resinous; bud
truly magnificent sizes. It is (still) heavily logged in scales triangular, obtuse, keeled at base, appressed,
clear cuts from natural stands including old growth slightly recurved at apex, shining chestnut brown,
(in this part of the world this means: forest that was persisting several years. Leaves spreading radially,
never cut before). Smaller sizes go to the paper in- directed obliquely forward, slightly incurved, esp. on
dustry, but big trees are prized for construction and coning shoots, 2.5–4.5(–5) cm long, ca. 1 mm wide,
special uses such as small aircraft, masts and spars narrowly linear, straight or curved, soft, rhombic to
for sailing ships, oars for rowing boats, ladders, broadly rhombic in cross-section, obliquely acute or
and sounding boards of musical instruments. Sitka acuminate at apex; amphistomatic, with 3–5 lines of
spruce has been widely used in plantation forestry stomata on each face; leaf colour dark green. Pollen
on poor acid soils in cool and wet climates such as cones axillary, 2–3 cm long, yellow. Seed cones ter-
the hills and moors of Ireland and Scotland; this minal, erect at first, soon pendulous, obliquely short
timber is used for pulp wood. In horticulture it finds pedunculate or nearly sessile, cylindro-conical or
less use; most plantings in large parks as specimen broad fusiform (when closed) often widest near base
trees date from the 19th century, and only a limited when opened, obtuse at apex, (8.5–)10–17(–18) cm
number of cultivars has been produced, mostly long, 4–6 cm wide with opened scales; colour (im-
dwarf forms. It requires a cool and moist climate. mature) green or purplish green, maturing to shiny
bright green, ripening to lustrous brown, old cones
dull grey-brown. Seed scales obovate-flabellate,
Picea smithiana (Wall.) Boiss., Fl. Orient. 5: 700. slightly convex, coriaceous, spreading very wide in
1884. Pinus smithiana Wall., Pl. Asiat. Rar. 3: 24, t. opened cones, 2–3 × 1.5–2.4 cm at mid-cone; abax-
246. 1832. Type: India: Himalaya, W. S. Webb & [?] ial surface finely striated, smooth, lustrous, often
Govan 6063 (lectotype C). Fig. 199, 200, 201 resinous; upper margin entire, rounded or slightly
obtuse, incurved; base cuneate. Bracts rudimenta-
Picea smithiana (Wall.) Boiss. var. nepalensis Franco, ry, ligulate, 4–5 mm long, entirely included. Seeds
Enum. Fl. Pl. Nepal 1: 26. 1978. ovoid-oblong, pointed at apex, 5–7 × 3–4 mm, dark
(red-) brown; seed wings ovate-oblong, 13–20 × 7–9 Picea spinulosa (Griff.) Beissn., Mitt. Deutsch.
mm, orange-brown. Dendrol. Ges. 15: 83. 1906. Abies spinulosa Griff.,
Itin. Notes: 259, 265, 275. 1848. Type: India: Sikkim,
Lachen River, Lachen, J. D. Hooker s.n., 1849 ([orig.
Distribution
mat. lost] neotype K, here designated).
Afghanistan: Hindu Kush; China: Xizang (Tibetan
Himalaya); W Himalaya: Himachal Pradesh, Kara- Picea spinulosa (Griff.) Beissn. var. yatungensis Silba,
koram, Kashmir Himalaya, Uttar Pradesh, Nepal Phytologia 68: 45. 1990.
TDWG codes: 34 AFG 36 CHT 40 NEP PAK WHM-
HP WHM-JK WHM-UT
Etymology
601
The species epithet means ‘somewhat thorny’ (Latin
Ecology
spinula = small thorn), but it is not clear what Grif-
Picea smithiana is a high mountain species, occur- fith had in mind with this.
ring in the Himalayas in a belt between (2300–)2500–
3600(–3750) m a.s.l. on alpine lithosols. The climate
Vernacular names
is moist monsoon, with abundant precipitation in
two rainy seasons, but becoming gradually drier in Sikkim spruce, East Himalayan spruce; Xizang yun-
the western parts of the range. It usually occurs with shan (Chinese)
Abies spectabilis, Pinus wallichiana and Tsuga dumo­
sa (eastern part of the range) and with Abies pindrow
Description
and Cedrus deodara in the western part. It is best
developed on northerly exposed slopes in Kashmir. Trees to 50–60 m tall, d.b.h. to 1.5–2.5 m; trunk
At lower elevations various broad-leaved trees, e.g. monopodial, straight; bark rough and scaly, on
Quercus spp., Acer spp., Prunus spp., Ulmus spp., large trunks fissured below, brownish grey or grey.
and Aesculus indica become dominant. Branches of first order long, spreading horizontally,
curved downward below, drooping at ends; branch-
es of second order long, slender, pendulous; crown
Conservation
pyramidal, in old trees broad columnar, domed, or
IUCN: LC open and irregular. Branchlets slender, flexible, soon
pendulous, pale yellowish grey or pale brown, prom-
inently ridged and deeply grooved, glabrous; pulvini
Uses
1–1.5 mm, narrow, at 60–80° from shoot. Vegetative
Idian or West Himalayan spruce is an important buds ovoid, obtuse, 5–7 mm long, not or only slightly
timber tree which can yield excellent construction resinous; bud scales ovate, obtuse, orange-brown or
material of large size. Its wood is used in interior red-brown, persisting several years. Leaves radially
house construction for roofs and floors. Large quan- spreading on pendulous shoots, on more horizontal
tities have been used as railway sleepers after treat- branches crowded above shoot, directed forward,
ment with preservatives to lengthen its durability parted below, 1.5–3.5 cm long, (0.7–)1–1.8(–2) mm
outdoors. Its relatively light weight combined with wide, narrowly linear, straight or slightly curved,
strength made it suitable for aircraft, in particular transversely rhombic to quadrangular in cross-sec-
gliders, but wood is no longer the main material for tion, keeled on both sides, pungent at apex; usually
these. Smaller sizes are now being pulped for pa- epistomatic, but sometimes 1–2 lines of stomata on
per, especially newsprint. Its long needles, droop- the upperside, 2 bands of 4–5 lines below (invers-
ing branches and bright green seed cones make it dorsal side); leaf colour lustrous dark green on the
an ornamental tree of considerable merit. It is not upperside, bluish white stomatal bands below. Pol-
very common in cultivation, perhaps due to prob- len cones axillary, 2–2.5 cm long, yellow when ripe.
lems with establishment and slow growth; it should Seed cones terminal, erect at first, pendulous at
be hardy in most countries with a cool temperate maturity, short pedunculate or sessile, cylindrical,
climate. narrowed at base, obtuse at apex, (4–)6–9(–12) cm
 long, (2.5–)3–4.5 cm wide with opened scales; col- Picea torano (Siebold ex K. Koch) Koehne,
our (immature) green or reddish green, ripening to Deutsche Dendrol.: 22. 1893. Abies torano Siebold
light reddish brown. Seed scales obovate or obtrul- ex K. Koch, Dendrol. 2 (2): 233. 1873. Type: Japan:
late, often irregular, 1.5–2 × 0.8–1.4 cm at mid-cone; [“Owari pr., Abies No. 1 torano ki”], P. F. von
abaxial surface finely striated, lustrous, undulate or Siebold s.n. sub Abies polita (lectotype L). Pl. 25
with irregular, longitudinal grooves, glabrous; upper
margin entire or denticulate; apex elongated, slight- Abies polita Siebold & Zucc., Fl. Japon. 2 (2): 20, t.
ly incurved, dentate; base cuneate. Bracts rudimen- 111. 1842 (nom. illeg., Art. 52.1); Picea polita (Siebold
tary, ligulate, 3–4 mm long, entirely included. Seeds & Zucc.) Carrière, Traité Gén. Conif.: 256. 1855.
ovoid-oblong, 3–5 mm, grey or brown; seed wings
602 obovate-oblong, 8–14 × 4–6 mm, orange-brown.
Etymology
The species epithet repeats one of the vernacular
Distribution
Japanese names of this species.
Eastern Himalaya: Bhutan and Sikkim (between the
Manas River and the Tista River), S Xizang [Tibet].
Vernacular names
TDWG codes: 36 cht 40 EHM-BH EHM-DJ EHM-SI
Tigertail spruce; Torano momi, Tora momi (Japa-
nese)
Ecology
This species occurs in the ‘cloud belt’ zone of the
Description
E Himalayas, at elevations between 2400 m and
3700 m a.s.l. (in S Xizang [Tibet] 2900 m to 3600 Trees to 30–40 m tall, d.b.h. to 1–1.3 m; trunk mo-
m). It grows on alpine lithosols. The climate is moist nopodial, straight; bark on trunk rough and scaly,
monsoon, with moderate to high annual precipita- flaking, fissured, pale grey, with pale brown inner
tion (1000 mm to 2000 mm), decreasing from south bark. Branches of first order long, spreading hori-
to north. It is usually associated with Pinus wallichi­ zontally or slightly ascending; branches of second
ana, Larix griffithiana, Abies densa, Tsuga dumosa, order relatively short, dense, spreading or assurgent;
Taxus baccata, and Juniperus wallichiana; at the up- crown broad pyramidal, domed or flat topped in
per forest limit dense growth of Rhododendron spp. old trees. Branchlets thick, very stout, shining yel-
replaces the conifers. At lower elevations various lowish brown, later grey, prominently ridged and
broad-leaved trees become more abundant: Acer deeply grooved, glabrous; pulvini strongly devel-
spp., Juglans regia, Fraxinus spp., Sorbus spp., Quer­ oped, 2–3 mm long, thickened at base, spreading at
cus pachyphylla, Prunus spp., and tree-like species of 70–90° from shoot. Vegetative buds ovoid or ovoid-
Rhododendron are the most common. oblong, obtuse or acute, 8–12 × 4–8 mm, not or only
slightly resinous; bud scales ovate, obtuse, smooth,
shining chestnut brown, appressed, persisting sev-
Conservation
eral years, leaving thick collars of imbricate perular
IUCN: LC scales around shoot bases. Leaves spreading radially,
curved, assurgent above shoot, very rigid, (1–)1.5–
2(–2.5) cm long, 1.8–2(–2.5) mm wide, linear, curved
Uses
or twisted, quadrate-rhombic in cross-section, very
No local uses have been recorded of this species, al- pungent to spinescent at apex; amphistomatic, with
though it is likely that its timber is being used for 4–6 lines of stomata on each face; leaf colour dark,
building houses. It is quite rare in cultivation despite lustrous green. Pollen cones axillary, 3–3.5 cm long,
considerable garden merit; it was introduced to reddish at first, yellow at maturity. Seed cones ter-
England from Sikkim in 1878. It seems to have failed minal, erect at first, pendulous at maturity, sessile,
from earlier sendings of seed via the botanic garden ovoid or ovoid-oblong, obtuse or tapering to a point
in Calcutta and is apparently not growing in Central (when closed) at apex, (5–)8–10(–12) cm long, 4–7 cm
Europe, where the winters may be too cold. wide with opened scales; colour (immature) green
or yellowish green, ripening to cinnamon brown
 603

1
2

5
3

Pl ate 2 5. Picea torano. 1. Habit of tree. 2. Branch with foliage. 3. Green seed cone. 4. Ripe seed cone.
5. Leaf. 6. Seed.
 or reddish brown. Seed scales cuneate-­obovate to something off the well beaten track seems to dem-
semi-orbicular (near base of cone), opening very onstrate itself most clearly.
wide, 2–2.8 × 1.5–2.5 cm at mid-cone; surface finely
striated, lustrous, glabrous; upper margin entire or
denticulate, rounded, convex, sometimes lacerate; Picea wilsonii Mast., Gard. Chron., ser. 3, 33: 133.
base cuneate. Bracts small, ligulate-linear, 6–7 mm 1903. Type: China: W Hubei, E. H. Wilson 1897
long, entirely included. Seeds ovoid, 5–7 × 3–4 mm, (holotype BM). Fig. 202
brown or grey-brown; seed wings obovate-oblong,
15–18 × 7–10 mm, orange-brown. Picea watsoniana Mast., J. Linn. Soc., Bot. 37: 419.
1906; Picea wilsonii Mast. var. watsoniana (Mast.)
604 Silba, Phytologia 68: 46. 1990.
Distribution
Picea wilsonii Mast. var. shanxiensis Silba, Phytolo-
Japan: Honshu, Kyushu, Shikoku. gia 68: 46. 1990.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Etymology
Ecology
This species was named after Ernest H. Wilson
Picea torano occurs in (low) mountains at eleva- (1876–1930), the famous plant hunter who travelled
tions between 400 m and 1850 m a.s.l., almost in- in China, Korea and Japan.
variably on podzolic soils of young volcanic rocks
such as lava flows and tuffs. The climate is cool,
Vernacular names
moist maritime, with annual precipitation exceed-
ing 1000 mm, the winters are cold and snowy, espe- Wilson spruce; Qinggian (Chinese)
cially at the higher elevations. There are some small
remnants of pure stands left, e.g. at the northern end
Description
of Lake Yamanaka, elsewhere it is mixed with Abies
homolepis, Larix kaempferi, Pinus densiflora and/or Trees to 40–50 m tall, d.b.h. to 1.3–1.5 m; trunk
broad-leaved trees, e.g. Betula, Fagus, Acer, Quercus mono­podial, straight; bark on trunk rough and
mongolica var. grosseserrata, Prunus maximowiczii, scaly, greyish brown or dark grey. Branches of first
and Zelkova serrata. order slender, spreading horizontally; branches of
second order short, slender, spreading, in some trees
becoming pendant; crown pyramidal and dense in
Conservation
young trees, columnar in old trees. Branchlets slen-
IUCN: LC der, thin, flexible, pale (yellowish) grey or buff grey,
finely ridged and grooved, glabrous; pulvini very
small, at 50–70° from shoot. Vegetative buds ovoid,
Uses
obtuse, 6–8 mm long, not resinous; bud scales ovate,
Tigertail spruce is uncommon and grows in inacces- obtuse, appressed, shining, dark brown or purplish
sible places, therefore it is not an important timber brown, persisting several years. Leaves spreading ra-
tree. In Japan it is a popular amenity tree, used in dially, above shoot more or less imbricate, directed
gardens and parks. Despite its striking foliage and forward, parted below, (0.8–)1–1.8(–2.2) cm long,
cones and conical habit in cultivation, it remains an 1–1.7 mm wide, linear, straight or slightly curved,
uncommon tree in gardens and arboreta in Europe mostly transversely rhombic in cross-section, obtuse
and is even less common elsewhere outside Japan. or acute at apex; stomata on all sides, 1–2 lines on
Despite its praise sung in earlier conifer handbooks each face above, 2–4 lines below; leaf colour glossy
(e.g. Dallimore & Jackson, 1966), a recent book on dark green. Pollen cones axillary, 2–3 cm long, yel-
cultivated conifers in the USA (Bitner, 2007) does lowish when ripe. Seed cones terminal, erect at first,
not even mention it. It is perfectly adapted to most later pendulous, very numerous, sessile or obliquely
climate zones in the temperate parts of the world short pedunculate, ovoid-oblong or cylindrical,
and should be grown much more often; it is there with obtuse apex, (4–)5–8 cm long, 2.5–3.5(–4) cm
where commercial conservatism and angst to try wide with opened scales; colour (immature) light
green or purplish green, ripening to pale brown or mate is continental, montane to subalpine, with low
dull reddish brown. Seed scales obovate-cuneate, precipitation and cold, long winters. In the montane
slightly convex, opening wide, 1.4–1.7 × 0.9–1.4 cm boreal coniferous forests of N Sichuan and S Gansu
at mid-cone; surface finely striated, in old cones it is usually growing with Picea asperata and P. pur­
nearly smooth, glabrous; upper margin entire or purea, also with P. meyeri. It is as a rule much rarer
erose-denticulate, rounded or truncate, sometimes than these. Betula albosinensis is the most common
obtuse; base cuneate. Bracts rudimentary, ligu- broad-leaved tree in these spruce forests.
late, 3–4 mm long, entirely included. Seeds ovoid,
pointed at apex, 3–4.5 × 2.5–3 mm, dark brown; seed
Conservation
wings obovate-oblong, 12–15 × 5–7 mm, yellowish or
light brown. IUCN: LC 605

Distribution Uses
China: Gansu, Hebei, Hubei, Shanxi, Shaanxi, W Si- Wilson spruce is an important timber tree in China
chuan. yielding wood for construction, railway sleepers,
TDWG codes: 36 CHC-HU CHC-SC CHN-GS CHN- mining props, carpentry, and furniture making, as
HB CHN-SA CHN-SX well as for pulp used in the paper (newsprint) in-
dustry. In horticulture, it is commonly planted as
an amenity tree in northern China and Russia and
Ecology
grows well in regions with long, steady winters, but
Picea wilsonii is a high mountain species occurring suffers from ‘late’ frosts in more maritime climates,
at elevations between 1400 m and 3000 m a.s.l. (max. especially in western Europe. In Europe and the
2100 m in the north of its range). The soils are most- USA it is seldom seen outside arboreta and botanic
ly non-calcareous, podzolic mountain soils. The cli- gardens.
 Pilgerodendron Florin, Svensk Bot. Tidskr. 24: 132. 1930. Type: Pilgerodendron
uviferum (D. Don) Florin [Juniperus uvifera D. Don] (Cupressaceae).

Named after Robert Pilger (1876–1953), German ing fibrous and fissured, exfoliating in long strips,
botanist at the Berlin Botanic Garden; Greek: den­ weathering grey-brown. Branches spreading hori-
dron = tree. zontally and assurgent at ends, higher order branches
slender, forming a narrowly conical to pyramidal, in
old trees broad rounded or even flat-topped crown.
Description
Foliage branches numerous, crowded, spreading or
606 See the species description. assurgent, ultimate branchlets short, slender but
rigid, densely covered by scale leaves and thereby
quadrangular, semi-deciduous. Leaves scale-like,
Distribution
decussate, occasionally in whorls of 3, short decur-
As for the species. rent at base, covering shoot, imbricate, forming 4(6)
rows, monomorphic, (broad) lanceolate, the free
part reflexed, narrowly triangular, carinate abaxially,
Taxonomic notes
2.5–6 × 1–2.2 mm on ultimate branchlets, on older
An overview of the taxonomic history and char- branchlets up to twice that size; margins entire; apex
acters of this monotypic genus has been given in obtuse; epistomatic, adaxial leaf face with numerous
A Monograph of Cupressaceae and Sciadopitys (Far- conspicuous stomata, leaf colour dark green, with a
jon, 2005a). Reasons to retain the genus and not whitish stomatal band. Pollen cones terminal, soli-
re-unite it with Libocedrus Endl., as suggested by tary, cylindrical, 5–10 × 2–2.5 mm; microsporophylls
a phylogenetic analysis based (mainly) on cpDNA 12–20, decussate, triangular to nearly rhombic, more
data (Gadek et al., 2000) were also outlined in that or less rostrate, subpeltate, bearing 4–8(–10) abaxial,
book, to which the interested reader is referred. small, globose pollen sacs. Seed cones terminal on
foliage branchlets, subtended by 4 decussate, nar-
rower and longer leaves, maturing within one year
Pilgerodendron uviferum (D. Don) Florin, Svensk to cones 8–12 × 4–6 mm, with distally spreading
Bot. Tidskr. 24: 133. 1930. Juniperus uvifera D. Don, scales. Bract-scale complexes in 2 decussate pairs,
in Lambert, Descr. Pinus, ed. 2, 2: 116. 1828; Libo­ elliptic to obtrullate, with upper (fertile) pair 7–11 ×
cedrus uvifera (D. Don) Pilg., in Engler & Prantl, 3–5 mm and lower pair lanceolate and half that size,
Nat. Pflanzenfam., ed. 2, 13: 389. 1926. Type: Chile: concavo-convex, with a large portion of the bract
Magallanes, Isla Madre de Dios, Puerto Molyneux, protruding subapically near the widest portion of
L. Savatier 140 (neotype K). Fig. 203, 204 the scale; margins papillose, colour reddish brown
or brown, the inner surface smooth, grooved or
ribbed towards base, lustrous light brown or whit-
Etymology
ish. Columella conspicuous, a small spike 2–3 mm
The species epithet means ‘bearing grapes’ (Latin long. Seeds 3–4 per cone, conical or triangular, with
uva = grape(s) or bunch of grapes). a basal hilum, 3 × 1.5–2 mm, pale yellowish brown;
wings 2, marginal, of very unequal size; largest wing
6–7 × 3–3.5 mm, obovate-elliptic; smallest wing a
Vernacular names
narrow strip.
Guaitecas Cypress; Ciprès de las Guaitecas (Chile),
Ten (Argentina), Ciprès (Spanish)
Distribution
S Argentina: Santa Cruz; S Chile: Aisén, Los Lagos,
Description
Magellanes.
Trees to 20 m tall, evergreen, dioecious; trunk mo- TDWG codes: 85 AGS-SC CLS-AI CLS-LL CLS-MG
nopodial, erect, to 1.5 m d.b.h. Bark brown, becom-
 spread clearance of the forests caused a considerable
Ecology
decline in this and other conifer species. Its value as
This species is a codominant tree along the edges of a timber source, although less than that of Fitzroya,
bogs and swamps of the coastal mountains, at alti- was still a sufficient factor to increase the pressure
tudes between 20 m and 750 m a.s.l. and growing on this tree, causing its virtual extinction in several
with Fitzroya cupressoides and the coniferous shrub areas. Continuing pressure until recently has caused
Lepidothamnus fonkii. It occurs also in large and this species to be listed in the Appendix I of CITES,
dense stands on wet mountain slopes of islands in effectively prohibiting the export of its wood.
shallow, water-logged acidic soil over metamorphic IUCN: VU (A1cd+2cd)
rocks with Fitzroya, Podocarpus nubigenus, Notho­
fagus nitida, Desfontainia spinosa, and the shrub 607
Uses
Tepualia stipularis, and with Nothofagus betuloides
and Drimys winteri in the southernmost parts of its The timber of this species has been used for build-
range. In more continental locations E of the crest of ing and construction of local farmsteads and other
the Andes it grows near mountain lakes. Resistance buildings in the past. The wood is decay resistant.
to light fires assists the regeneration of this shade in- There is still local use and trade but, due to dem-
tolerant species, but intense fires will and have killed inished resources, much less than in the past and
many trees. The climate is cool maritime with ex- export is now prohibited. Introduced to England by
tremely high precipitation in the coastal ranges. William Lobb in 1849, it is still uncommon in culti-
vation and almost restricted to arboreta and botanic
gardens, which is unfortunate because it grows into
Conservation
a handsome, conical shrub and is perfectly hardy
During the period of colonization by Europeans, and suitable on acidic soils in all countries with a
when agriculture was introduced on a large scale in relatively mild winter.
many lowland and colline regions of Chile, wide-
 Pinus L., Sp. Pl. 2: 1000. 1753. Type: Pinus sylvestris L. (Pinaceae).

Pinea Wolf, Gen. Pl.: 156. 1776. Type: Pinus pinea serotinous, (obliquely) ovoid to cylindrical, 2–60
L. Strobus Opiz, Lotos 4: 94. 1854. Type: Strobus cm long. Bracts conspicuous at pollination stage,
weymouthiana Opiz [Pinus strobus L.]. Caryopitys not growing with seed scales. Seed scales persistent,
Small, Fl. SE United States: 29. 1903. Type: Caryo­ obovate to oblong, thin or thick woody, attached
pitys edulis (Engelm.) Small [Pinus edulis Engelm.]. spirally to a slender or thick rachis; exposed portion
Apinus Neck. ex Rydb., Bull. Torrey Bot. Club 32: (apophysis) variously thickened and/or elongat-
597. 1905. Type: Apinus cembra Neck. ex Rydb. [Pi­ ed, with a terminal or dorsal umbo, which may be
608 nus cembra L.]. Leucopitys Nieuwl., Amer. Midl. armed with a spine or prickle. Seeds usually slightly
Naturalist 3: 69. 1913 (nom. illeg.). Type: Leucopitys flattened, with an adnate or articulate membranous
strobus (L.) Nieuwl. [Pinus strobus L.]. Ducampo­ wing derived from adaxial part of seed scale and
pinus A. Chev., Rev. Bot. Appliq. 24: 30. 1944. Type: several times larger than seed or reduced. Seedlings
Ducampopinus krempfii (Lecomte) A. Chev. [Pinus with 3–24 cotyledons.
krempfii Lecomte]
113 species.
Pinus is the classical Latin name for pines.
Distribution
Description
North America: from Yukon to Newfoundland
Evergreen monoecious trees, less commonly shrubs; and from SE Alaska and British Columbia south
resin canals in wood, bark, leaves and often cones. into Mexico and Central America as far as Nica-
Trunk of trees monopodial, branching in pseudo- ragua; in E and SE USA to Florida; Bahamas, Car-
whorls, branches assurgent and repeating prima- ribean (Cuba, Hispaniola). Eurasia (N Africa): Atlas
ry branching arrangement (Rauh’s model). Bark Mountains of Morocco and Algeria; from the Ibe-
furrowed or plated, to thin and scaly or thin and rian ­Peninsula through S and Central Europe to
smooth. Shoots dimorphic, with long shoots and E Europe and Turkey, Caucasus, Syria and Lebanon;
dwarf shoots. Primary leaves (cataphylls) scale-like, Scotland (disjunct); from Scandinavia through Rus-
enclosing “winter buds” (primordial long shoots sia and Siberia to Kamchatka and Sachalin; Sino-
or ovuliferous strobili), subtending dwarf shoot Himalayan mountain system, China, Japan, Taiwan,
buds or pollen strobili, early deciduous or persist- Indochina to N Sumatera; Philippines.
ent. Secondary leaves (needles) borne in fascicles of
(1–)2–5(–8) on dwarf shoots; fascicles surrounded at
Synopsis
base by an early deciduous or persistent sheath of
bud scales or their remnants, persisting 2–30 years The genus Pinus is the only conifer genus for which
and falling as fascicles; length of needles 2.5–50 cm, several comprehensive studies investigating phylo-
width 0.5–2.5(–7) mm, acicular (one species lan- genetic relationships, based on a number of different
ceolate), plano-convex or triangular (rarely terete gene sequences, have been published. There is also
or flat) in cross-section, entire or serrulate, epi­ comprehensive work available on comparative mor-
stomatic or amphistomatic (one species occasionally phology, including some cladistic analyses. Conse-
hypostomatic). Pollen cones spirally arranged near quently, it is possible to set up a classification that is
proximal end of new long shoots, ovoid-oblong to informed by (if not entirely based on) a hypothesis
cylindrical; consisting of a thin axis with numerous of phylogenetic relationships. The most important
spirally arranged, (sub-)peltate microsporophylls, and consistent result of these phylogenetic analyses
each bearing two pollen sacs containing bisaccate is the confirmation that the genus naturally divides
pollen. Seed cones subterminal, borne singly or into two subgenera. This leaves all earlier classifica-
more commonly clustered, pedunculate, maturing tions that recognized more than two primary divi-
in second year, or rarely in third year, shed early or sions as essentially artificial; the result of a choice
variously persistent, initially erect; mature cones of characters rather than a reflection of evolution-
pendulous or spreading, opening soon or variously ary relationships. The first comprehensive formal
classification informed primarily by phylogeny was P. luchuensis, P. taiwanensis, P. densi­
published by Price et al. in Richardson (1998). The flora, P. sylvestris, P. resinosa, P. mugo,
second was given in my book Pines (Farjon, 2005b), P. uncinata, P. massoniana, P. tropica­
with a review of the more recent results of molecular lis, P. nigra
phylogenetic analyses. The latter classification, with Section Trifolius Duhamel
a few minor amendments, is adopted here. Subsection Contortae Little & Critchfield
Species: Pinus contorta, P. banksiana,
Genus Pinus L. P. virginiana, P. clausa
Subsection Ponderosae Loudon
Subgenus Strobus Lemmon Species: Pinus ponderosa, P. ari­
Section Parrya Mayr zonica, P. jeffreyi, P. engelmannii, 609
Subsection Nelsoniae Van der Burgh P. coulteri, P. sabiniana, P. torreyana,
Species: Pinus nelsonii P. hartwegii, P. montezumae, P. devo­
Subsection Balfourianae Engelm. niana, P. pseudostrobus, P. douglasi­
Species: Pinus balfouriana, P. arista­ ana, P. maximinoi, P. durangensis
ta, P. longaeva Subsection Australes Loudon
Subsection Rzedowskianae Carvajal Species: Pinus palustris, P. carib­
Species: Pinus rzedowskii, P. maxi­ aea, P. elliottii, P. glabra, P. cubensis,
martinezii, P. pinceana P. occidentalis, P. pungens, P. rigida,
Subsection Cembroides Engelm. P. serotina, P. taeda, P. leiophylla,
Species: Pinus cembroides, P. culmini­ P. lumholtzii, P. herrerae, P. echinata,
cola, P. edulis, P. remota, P. mono­ P. lawsonii, P. pringlei, P. teocote,
phylla, P. quadrifolia P. patula, P. greggii, P. attenuata,
Section Quinquefolius Duhamel P. muricata, P. radiata, P. jaliscana,
Subsection Gerardianae Loudon P. tecunumanii, P. oocarpa, P. luz­
Species: Pinus gerardiana, P. bunge­ mariae, P. praetermissa
ana, P. squamata
Subsection Krempfianae Little & Critch-
Keys for the genus Pinus
field
Species: Pinus krempfii The keys for the genus Pinus follow the taxonomic
Subsection Strobi Loudon classification presented above. Keys based on taxo-
Species: Pinus strobus, P. albicaulis, nomic classifications as opposed to those based on
P. flexilis, P. strobiformis, P. ayaca­ geographic groupings have the advantage that an in-
huite, P. monticola, P. lambertiana, dividual plant growing outside its native range can
P. cembra, P. sibirica, P. peuce, P. wal­ still be identified with them when the provenance
lichiana, P. bhutanica, P. armandii, of that plant is unknown. The genus Pinus has prob-
P. amamiana, P. koraiensis, P. pumila, ably more species planted outside their natural
P. parviflora, P. morrisonicola, P. fen­ ranges than any other genus in conifers. Excluded
zeliana, P. wangii, P. dalatensis from the keys are the four hybrid species Pinus
Subgenus Pinus ×densithunbergii, P. ×hakkodensis, P. ×neilreichiana
Section Pinus and P. ×rhaetica.
Subsection Pinaster Mayr ex Koehne
Species: Pinus pinaster, P. held­reichii,
Key to the subgenera of Pinus
P. brutia, P. halepensis, P. pinea,
P. ­canariensis, P. roxburghii, P. latteri, 1a. Pulvini decurrent. Leaves with 2 vascular bun-
P. merkusii dles and variously positioned resin ducts, with
Subsection Pinus (syn. Subsection Sylves­ a persistent (rarely deciduous) fascicle sheath.
tres Loudon) Seeds with an articulate, rarely adnate, wing
Species: Pinus densata, P. yunnanen­ Subgen. Pinus
sis, P. kesiya, P. thunbergii, P. tabuli­ 1b. Pulvini not decurrent. Leaves with a single
formis, P. henryi, P. hwangshanensis, vascular bundle and external (marginal) resin
 ducts, with a deciduous (rarely persistent) fas- sometimes medial or septal but very rarely 1–2
cicle sheath. Seeds with an adnate, rarely artic- ducts external. All New World species
ulate, wing or wingless when free from the seed Subsect. Australes
scale Subgen. Strobus
Note: There is in fact much similarity (and vari-
ation) among the species of the latter two subsec-
Key to the subsections of Subgenus Pinus
tions and many species cannot be reliably assigned
1a. Shoots uni-nodal; buds not resinous. Leaves in to their respective subsections on the basis of their
fascicles of only 2 or only 3 (rarely 2 plus some morphology. These subsections, while supported by
3 in P. brutia). Seed cones 6–20(–25) cm long; phylogenetic analyses based on DNA sequence data,
610 seed scales thick woody, rigid (except in P. held­ apparently contain much homoplasy (parallel evolu-
reichii); apophyses of seed scales lustrous brown tion or convergence) in their morphological charac-
(except in P. heldreichii); umbo unarmed or at ter states. It is therefore advised to try both keys to
most with a minute, deciduous prickle (except identify a species that belongs to any one of these
spinaceous in P. pinaster). All Old World spe- two subsections.
cies Subsect. Pinaster
1b. Not this combination of character states 2
Key to the species of Pinus Subsection
2a. Buds (slightly) resinous or rarely not resinous.
Australes
Leaves in fascicles of only 2 or 2 and 3 (only 3
in P. kesiya). Seed cones deciduous or persist- 1a. Scales of leaf fascicles deciduous, leaving the
ent for a few years but always opening soon, fascicles without a sheath before falling 2
2–10(–12) cm long; seed scales thin woody, rig- 1b. Scales of leaf fascicles persistent, retaining a
id; umbo unarmed or with a minute, deciduous sheath at the base of the fascicles when falling
or sometimes persistent prickle. All but 2 Old 3
World species Subsect. Pinus 2a. Leaves in fascicles of 3 (exceptionally 2 or 4),
2b. Not this combibation of character states 3 (15–)20–30(–40+) cm long, pendulous
3a. Bark on lower trunk of trees with small plates P. lumholtzii
or flakes. Shoots uni-nodal. Fascicle sheaths 2b. Leaves in fascicles of (2–)3–5(–6), (4–)6–15(–
shorter than 10 mm; leaves in fascicles of only 17) cm long, rigid or pliant, not pendulous
2, 2–8(–10) cm long, rigid, curved or twisted; P. leiophylla
resin ducts in leaves medial. Seed cones (semi-) 3a. New foliage shoots multi-nodal (producing
persistent, (2–)3–8 cm long, more or less seroti- more than 1 whorl of branches or buds in a sin-
nous or sometimes opening soon in warm sun- gle growing season) 4
shine; seed scales thin woody, rigid. All New 3b. New foliage shoots uni-nodal 10
World species Subsect. Contortae 4a. Seed cones deciduous or semi-persistent, open-
3b. Not this combination of character states 4 ing soon 5
4a. Leaves in fascicles of 3 or 5, less commonly on 4b. Seed cones persistent, serotinous 7
the same tree 2, 4 or 6(–8), rarely only 2 (P. pon­ 5a. Leaves in fascicles of 2, some times of 3; fascicle
derosa of northern populations), straight (not sheaths of new leaves 9–12 mm long P. cubensis
rigidly curved, contorted or twisted); fascicle 5b. Leaves in fascicles of (2–)3–4(–5); fascicle
sheaths of new leaf fascicles at least 15 mm long, sheaths of new leaves 15–30 mm long 6
usually 20–35(–40) mm long; resin ducts in 6a. Buds resinous. Leaves rigid, (1.2–)1.4–1.8 mm
leaves medial and sometimes 1–2 internal. Seed wide P. caribaea
cone scales opening soon or sometimes more 6b. Buds not resinous. Leaves lax, drooping, 0.7–
slowly (cones not serotinous). All New World 0.9(–1) mm wide P. patula
species Subsect. Ponderosae 7a. Leaves in fascicles of only 2, thick and rigid,
4b. Not this combination of character states 5 1.3–2 mm wide. Seed cones 5–7(–8) cm long;
5. Leaves in fascicles of 2–3, 3 only, or 3, 4 and 5, umbos of seed scales with a curved spine or
not of 5 only (or more than 5) and only P. gla­ sharp, persistent prickle P. muricata
bra and P. muricata with fascicles of 2 only; leaf 7b. Leaves in fascicles of 2 and 3, only 3, or rarely 4
resin ducts usually or predominantly internal, or 5, rigid, 1–1.6 mm wide. Seed cones (5–)7–15
 cm long; umbos of seed scales unarmed or with umbo of seed scales unarmed or with a minute,
a minute, deciduous prickle 8 deciduous prickle 16
8a. Apophyses of seed scales flat or slightly raised, 15b. Seed cones ovoid-conical, ovoid-attenuate,
more or less equally on all sides of the cone; narrowly conical or oblong when closed, with
umbo with a minute, deciduous prickle a short (maximum 20 mm long) peduncle
P. greggii or nearly sessile; umbo of seed scales with a
8b. Apophyses of seed scales prominently to ex- minute, deciduous or strong and sharp prickle
tremely raised at least on one side of the cone or spine 19
(in most cones); umbo unarmed 9 16a. Leaves in fascicles of 3, rarely of 4, rigid, 1.2–1.6
9a. Buds resinous. Pollen cones yellowish before mm wide P. luzmariae
anthesis. Seed cones ovoid-oblong to ovoid- 16b. Leaves in fascicles of 3, 4 or 5, pliant, lax and/or 611
attenuate P. attenuata drooping (sometimes more rigid), 0.5–1.6 mm
9b. Buds not resinous. Pollen cones pink to reddish wide 17
before anthesis. Seed cones ovoid, asymmetri- 17a. Seed cones deciduous, leaving basal scales on
cal in most cones, or nearly symmetrical if the the peduncle not falling with the cone
apophyses are only slightly raised P. radiata P. praetermissa
10a. Leaves (3–)4–12(–15) cm long, in fascicles of 17b. Seed cones persistent or deciduous, if decidu-
only 2 or 2 and 3. Buds (slightly) resinous 11 ous, falling intact with the peduncle attached to
10b. Leaves (7–)10–25(–45) cm long, in fascicles of the cone base 18
2 and 3, sometimes 4 or 5, or only 3 (if 2 and 3 18a. Seed cones deciduous, opening soon; apo­
or only 3, usually longer than 12 cm). Buds not physes of the seed scales raised P. tecunumanii
resinous or resinous (if resinous, leaves usually 18b. Seed cones persistent, semi-serotinous, apo-
longer than 12 cm) 14 physes of seed scales nearly flat or slightly
11a. Leaves in fascicles of only 2, slender, 0.7–1.2 raised P. oocarpa
mm wide. Seed cones solitary or in pairs 19a. Leaves lax and drooping, very slender, (0.5–)
P. glabra 0.6–0.8 mm wide P. jaliscana
11b. Leaves in fascicles of 2, 3 or sometimes 4, ro- 19b. Leaves rigid or pliant, 1–1.7 mm wide 20
bust, 1–1.5(–2) mm wide. Seed cones solitary or 20a. Buds (slightly) resinous. Seed cone scales thin;
more commonly in whorls of 2–6 12 umbo of scales armed with a prickle or spine
12a. Seed cones serotinous; seed scales with promi- 21
nently raised apophyses ending in a hard, sharp 20b. Buds not resinous. Seed cone scales thick;
spine P. pungens umbo of scales unarmed or with a minute, de-
12b. Seed cones opening soon or serotinous; seed ciduous prickle 23
scales with slightly raised apophyses ending in 21a. Leaves in fascicles of 3–5; fascicle sheaths of
a small but persistent prickle 13 new leaves (8–)10–15 mm long. Seed cones (lus-
13a. Seed cones persistent, opening soon or seroti- trous) dark brown P. occidentalis
nous, ovoid-conical when closed; umbos of 21b. Leaves in fascicles of (2–)3, occasionally 4; fas-
seed scales with a slender, curved prickle. Seed cicle sheaths of new leaves 15–25 mm long. Seed
wings 3–4 times longer than the seeds P. rigida cones light brown 22
13b. Seed cones semi-persistent, opening soon, nar- 22a. Leaves in fascicles of 2 and (more numerous) 3.
rowly ovoid or ovoid-oblong when closed; um- Seed cones deciduous, opening soon P. taeda
bos of seed scales with a short, stout prickle. 22b. Leaves in fascicles of 3, occasionally 4. Seed
Seed wings 2–3 times longer than the seeds cones persistent, opening soon or serotinous
P. echinata P. serotina
14a. Leaves only in fascicles of 3. Seed cones small 23a. Leaves 20–45 cm long, in fascicles of 3, rarely 2
(2–)3–3.5(–4) cm long, deciduous P. herrerae or 5. Pollen cones 4–7(–8) cm long. Seed cones
14b. Leaves in fascicles of (2–)3, 4 or 5. Seed cones (15–)20–25 cm long P. palustris
usually longer than 4 cm, deciduous or persist- 23b. Leaves (7–)10–25(–30) cm long, in fascicles
ent 15 of 2, 3, 4 or 5. Pollen cones 1–4 cm long. Seed
15a. Seed cones broadly ovoid to subglobose when cones (3–)4–15(–18) cm long 24
closed, with a (15–)20–35 mm long peduncle;
 24a. Leaves in fascicles of 2 and 3. Pollen cones
Key to the species of Pinus Subsection
purplish red before anthesis. Seed cones (7–)
Pinaster
9–15(–18) cm long; apophyses of seed scales
raised. Seeds 6–7 mm long, with a 20–30 mm The species in this subsection are naturally occur-
long wing P. elliottii ring in the Canary Islands, the Mediterranean, the
24b. Leaves in fascicles of 3–5 (very rarely 2). Pollen western Himalaya and tropical Southeast Asia. They
cones yellowish green or yellow before anthesis. are likely the remnant species of a past distribution
Seed cones (3–)4–8(–10) cm long; apophyses of of numerous pines that bordered the northern coasts
seed scales nearly flat to slightly raised. Seeds of the Thetis Ocean from the (early?) Cretaceous to
3–6 mm long, with a 12–18 mm long wing 25 the Miocene (Klaus, 1988). With one exception, they
612 25a. Bark on the trunk breaking into small plates. have seed cones with hard, rigid scales of which the
Seed cones persistent, semi-serotinous; apo- apophyses are lustrous brown with unarmed umbos.
physes of seed scales lustrous brown P. pringlei The exception, Pinus heldreichii, was formerly clas-
25b. Bark on the trunk deeply fissured. Seed cones sified as closely related to Pinus nigra and producing
deciduous, opening soon; apophyses of seed hybrids with it (see e.g. Vidaković, 1991), but phylo-
scales light brown 26 gentic analyses using DNA sequence data have dem-
26a. Leaves (7–)10–15(–18) cm long, mostly in fasci- onstrated that these two species end up in different
cles of 3, but some of 2, 4 or 5, straight or curved. clades, with P. heldreichii closely related (sister to)
Seed cones (3–)4–6(–7) cm long; umbos of seed the species here classified in subsection Pinaster and
scales with a minute, deciduous prickle P. nigra positioned within the clade forming subsec-
P. teocote tion Pinus (Wang et al., 1999; Geada López et al.,
26b. Leaves 12–20(–25) cm long, mostly in fascicles 2002). For this reason, P. heldreichii is here classified
of 3 and 4, sometimes of 5 (very rarely of 2), in subsection Pinaster.
straight. Seed cones 5–8(–9) cm long; umbos of
seed scales unarmed P. lawsonii 1a. Leaves in fascicles of only 3, 20–30(–35) cm
long, pliant and drooping 2
1b. Leaves in fascicles of only 2 or 2 and sometimes
Key to the species of Pinus Subsection
3, 6–25(–27) cm long, rigid or in one species
Contortae
pliant but not drooping 3
1a. Leaves curved, (0.7–)1.5–2.5(–3) mm wide. 2a. Fascicle sheaths of new leaves 25–30 mm long.
Seed cones solitary or in whorls of 2–5. Seed Seed cones persistent, opening slowly; apophy-
wings 10–12 mm long 2 ses of scales very strongly developed, conical
1b. Leaves straight but twisted, 1–1.5 mm wide. and often recurved P. roxburghii
Seed cones solitary or in pairs. Seed wings 15– 2b. Fascicle sheaths of new leaves 15–20 mm long.
20 mm long 3 Seed cones deciduous, opening soon; apophy-
2a. Pollen cones orange-red before anthesis. Seed ses of scales prominently raised but not conical
cones ovoid but asymmetrical at base when (except sometimes in some basal scales)
closed, opening slowly; umbos of seed scales P. canariensis
with a variable, persistent prickle P. contorta 3a. Fascicle sheaths of new leaves 20–30 mm long;
2b. Pollen cones yellow before anthesis. Seed cones leaves thick, 1.5–2 mm wide. Seed cones 10–
asymmetric, curved when closed, serotinous; 20(–22) cm long, persistent, opening slowly;
umbos of seed scales unarmed P. banksiana umbo on scales forming a central, rigid spine
3a. Seed cones conical, slightly oblique when P. pinaster
closed; apophyses of seed scales prominently 3b. Fascicle sheaths of new leaves shorter than 20
raised; umbos with a strong, slender prickle to mm; leaves 0.7–1.5(–1.8) mm wide. Seed cones
5 mm long P. virginiana shorter than 13(–15) cm, deciduous or persist-
3b. Seed cones narrowly ovoid when closed; apo- ent; umbos unarmed or with a minute prickle
physes of seed scales slightly raised; umbos un- 4
armed or with a small prickle P. clausa 4a. Seed scales of cones thin and flexible; apophy-
ses of scales dull brown; umbos with a minute
prickle. Leaves usually curved P. heldreichii
4b. Seed scales of cones thick and rigid; apophy- tree, ­especially those native to Mexico, but also from
ses of scales lustrous brown; umbos unarmed. elsewhere. Fascicles with 2–3 are common, as are
Leaves straight and/or twisted 5 3–5, either with predominance at the lower number,
5a. Bark on trunk breaking into large plates. Fas- or at the higher, or sometimes more equally distrib-
cicle sheaths of new leaves short, to ca. 10 mm uted. Rarely does the number exceed 5, but in a few
long. Seed cones broadly ovoid to subglobose species in Mexico this sometimes occurs and it can
when closed. Seeds 15–20(–22) mm long, thick, run up to 8. Several species never have more than
with a rudimentary wing much shorter than 2 and several others always have 5, but those with
the seed P. pinea 3 are often more likely to have some variation, ei-
5b. Bark on trunk breaking into small or elongated ther downward or upward. It is therefore necessary
plates. Fascicle sheaths of new leaves (9–)10–20 to count more than a few fascicles in many cases, 613
mm long. Seed cones ovoid-conical or oblong- if possible from different parts of the tree or, in a
conical when closed. Seeds less than 10 mm population, from a few trees. The number of leaves
long, with a wing at least 2 × the length of the in a fascicle is not a sufficient diagnostic character
seed 6 of species and needs to be supplemented with other
6a. Seed cones persistent, serotinous; apophyses of information.
scales nearly flat or slightly raised. Resin ducts
in leaves external (marginal). Seedlings without 1a. Leaves in fascicles of only 2 2
a ‘grass stage’ 7 1b. Leaves in fascicles of 2 or 3, or only 3 12
6b. Seed cones deciduous, opening soon; apophy- 2a. Shrubs or small trees. Leaves (2.3–)3–7(–8) cm
ses of scales prominently raised. Resin ducts in long, rigid and curved. Seed cones semi-per-
leaves medial. Seedlings commonly developing sistent, obliquely ovoid when closed; apophyses
a ‘grass stage’ 8 of scales on one side of the cone (prominently)
7a. Leaves in fascicles of only 2, (6–)7–10(–12) cm raised 3
long, 0.7–0.8 mm wide P. halepensis 2b. Trees. Leaves usually longer than 7 cm, straight
7b. Leaves in fascicles of 2 or sometimes 3, (5–)10– or when curved often pliant. Seed cones either
18(–29) cm long, 1–1.5 mm wide P. brutia deciduous or long persistent (rarely semi-per-
8a. Leaves 15–20 cm long, 1 mm wide, pliant. Seed sistent), more or less symmetrical when closed;
wings ca. 4 × as long as the seeds P. merkusii apophyses of scales flat or raised on all sides of
8b. Leaves 15–25(–27) cm long, 1.5 mm wide, rigid. the cone 4
Seed wings not more than 3 × as long as the 3a. Shrubs, rarely small trees. Apophyses of seed
seeds P. latteri scales nearly flat to prominently raised P. mugo
3b. Small, monopodial trees. Apophyses of seed
scales all raised, most prominently on one side
Key to the species of Pinus Subsection Pinus
of the cone, curving backward P. uncinata
Counting leaves in fascicles requires a few guide- 4a. Bark on trunk with deep, longitudinal fissures.
lines to make sure one gets the correct information Umbo of seed cone scales unarmed (without a
from it. The adult type needle leaves of pines grow spine or prickle) P. thunbergii
from dwarf shoots and are held in a fascicle sheath 4b. Bark on trunk breaking into large and/or ir-
formed of bud scales. In the subgenus Pinus these regular plates. Umbo of seed cone scales with at
sheaths are mostly persistent (two exceptions) and least a minute (but possibly deciduous) prickle
in the subgenus Strobus they are mostly decidu- 5
ous (one exception). Leaves may fall off singularly 5a. Fascicle sheaths of new leaves 15–20 mm long;
from older fascicles with deciduous sheaths; for leaves brittle (breaking easily). Pollen cones red
that reason it is necessary to count leaves only on or purple before anthesis P. resinosa
recent cohorts of leaves with more or less intact 5b. Fascicle sheaths of new leaves 5–15 mm long;
fascicle sheaths. Needle leaves of pines are grouped leaves rigid or pliant (not breaking easily). Pol-
in fascicles of 1–8. There is only one species with len cones yellow or at most tinged red 6
consistently single leaves (P. monophylla); common 6a. Leaves thick, robust, 1–2 mm wide 7
numbers are 2, 3 and 5. However, there are numer- 6b. Leaves thin, slender, 0.6–1 mm wide 8
ous species in which the numbers vary on a single
 7a. Leaves 4–7(–12) cm long; resin ducts in leaves 14b. Bark on trunk with irregular fissures. Seed
external (marginal). Seed cones (2–)3–7 cm cones when closed narrowly ovoid, 4–6 cm
long, dull light brown. Seeds 3–5 mm long with long. Seeds 4–6 mm long P. densata
a 10–15 mm long wing P. sylvestris 15a. Leaves (15–)20–30 cm long, 1.5 mm wide,
7b. Leaves (4–)8–16(–18) cm long; resin ducts in straight and rigid. Umbo of cone scales un-
leaves medial. Seed cones (3.5–)5-10(–12) cm armed P. tropicalis
long, slightly lustrous light brown. Seeds (4–) 15b. Leaves 7–20(–30) cm long, 1–1.2 mm wide,
6–8 mm long with a 15-25 mm long wing straight, contorted or drooping, pliant. Umbo
P. nigra of cone scales mucronate, or at least with a
8a. Leaves ca. 1 mm wide, rigid, straight. Seed short prickle 16
614 cones deciduous; apophyses of seed scales dull 16a. Bark on trunk with long, vertical fissures. Fas-
brown P. densiflora cicle sheath of new leaves 15–20 mm long. Seed
8b. Leaves 0.6–1 mm wide, pliant, straight or cones deciduous P. massoniana
curved. Seed cones persistent; apophyses of 16b. Bark on trunk with irregular plates. Fasci-
seed scales lustrous brown 9 cle sheath of new leaves 10–15 mm long. Seed
9a. Seed cones 2.5–5 cm long, ovoid when closed. cones persistent P. yunnanensis
Wings of seeds 9–12 mm long. Leaves 7–12 cm
long P. henryi
Key to the species of Pinus Subsection
9b. Seed cones 3–6(–8) cm long, narrowly ovoid
Ponderosae
when closed. Wings of seeds 10-20 mm long.
Leaves (5–)10–17 cm long 10 The species in this subsection range from western
10a. Fascicle sheaths of new leaves 5–10 mm long. North America into Mexico and Central America.
Apophyses of seed scales nearly flat; umbo de- The group consists of a very widespread species,
pressed with a short, persistent prickle P. ponderosa, some Tertiary relicts now confined to
P. hwangshanensis the two Californias, and taxa that become increas-
10b. Fascicle sheaths of new leaves 10–14 mm long. ingly diverse genetically and morphologically in
Apophyses of seed scales mostly slightly raised; Mexico and Central America, where they are most
umbo flat or pyramidal, with a minute, decidu- likely still evolving. The relicts are most distinct,
ous prickle or unarmed 11 while the evolving species are mostly variable and
11a. Pollen cones yellow. Umbo of seed scales nearly can be difficult to separate on the characters used
flat, with a minute, deciduous prickle or un- in this key. Reference to the full descriptions is here
armed. Seed wings 15–20 mm long most necessary to arrive at a determination with
P. taiwanensis confidence. For counting numbers of leaves per fas-
11b. Pollen cones tinged red. Umbo of seed scales cicle see comments under the key to the species of
slightly raised, armed with a sharp, persistent Pinus Subsect. Pinus.
prickle. Seed wings 10–15 mm long
P. luchuensis 1a. Leaves in fascicles of not more than 3, either
12a. New foliage shoots multi-nodal (producing 2–3 or only 3 2
more than 1 whorl of branches or buds in a sin- 1b. Leaves in fascicles of 3, 4, or 5, rarely 2 or more
gle growing season). Leaves in fascicles of only than 5 5
3, long and thin, lax and drooping P. kesiya 2a. Seed cones deciduous, opening soon; scales
12b. New foliage shoots uni-nodal. Leaves in fas- thin woody; apophyses raised but not elongat-
cicles of 2 or 3, short or long but not lax and ed into a strongly curved spine 3
drooping 13 2b. Seed cones persistent, opening slowly; scales
13a. Buds slightly resinous. Leaves 6–15 cm long, thick woody; apophyses extremely raised and
rigid 14 elongated into a strongly curved spine 4
13b. Buds not resinous. Leaves 7–30 cm long, rigid 3a. Buds resinous. Pollen cones red or magenta be-
or pliant 15 fore anthesis. Seed cones 5–12 cm long
14a. Bark on trunk breaking into large plates. Seed P. ponderosa
cones when closed ovoid, 5–8.5 cm long. Seeds
6–8 mm long P. tabuliformis
3b. Buds not resinous. Pollen cones yellow or with 12a. Seed cones 7–10 cm long; umbo unarmed.
a purplish tinge. Seed cones (10–)12–25(–30) Seeds 4–5 mm long P. douglasiana
cm long P. jeffreyi 12b. Seed cones 7–20 cm long; umbo obtuse but
4a. New foliage shoots multi-nodal (producing strongly developed, or with a small, persistent
more than 1 whorl of branches or buds in a prickle. Seeds 5–7 mm long 13
single growing season). Leaves thick and rig- 13a. Bark on trunk breaking into small plates, not
id, 1.9–2.2 mm wide. Seed cones solitary or in with deep longitudinal fissures. Seed cones soli-
whorls of 2–5 P. coulteri tary or more often in whorls of 3–6; apophyses
4b. New foliage shoots uni-nodal. Leaves slender, flat or moderately raised P. montezumae
pliant, 1.5 mm wide. Seed cones solitary, rarely 13b. Bark on trunk breaking into elongated plates
in pairs P. sabiniana and deep, longitudinal fissures. Seed cones soli- 615
5a. Leaves in fascicles of 3–5, rarely 2 6 tary or in whorls of 2–3; apohyses very variable,
5b. Leaves in fascicles of 5, rarely 3, 4, or 6, very from nearly flat to extremely raised or conical,
rarely 8 7 especially towards the base of the cone
6a. Leaves in fascicles of (2–)3(–4), rarely 5, 20–35 P. pseudostrobus
cm long, 1.5–2 mm wide P. engelmannii
6b. Leaves in fascicles of 3–5, 10–20(–25) cm long,
Key to the subsections (and some species) of
0.9–1.8 mm wide P. arizonica
Pinus Subgenus Strobus
7a. Leaves thick, rigid, ca. 2 mm wide. Pollen cones
before anthesis yellow, not tinged pink, purplish The morphological characters of species in this
or brown. Seed cones persistent, opening slow- subgenus are highly diverse but also demonstrate
ly, broadly ovoid when closed P. torreyana convergent evolution of certain traits, which means
7b. Leaves slender, rigid or pliant, 0.6–1.6 mm that similar character states do appear in phylo­
wide. Pollen cones before anthesis usually genetically not closely related species. In the classi-
tinged pink, purplish or brown. Seed cones de- fication adopted here (see Farjon, 2005b: 218–224)
ciduous, opening soon, ovoid-oblong or ovoid- some weight has been given to certain autapo­
attenuate when closed 8 morphies (characters unique in a species or lineage
8a. Leaves (6–)10–20(–24) cm long 9 that do not inform us about their relationships). In
8b. Leaves (15–)20–40(–45) cm long 10 a few cases the subsections therefore may diverge
9a. Leaves relatively thick, rigid, (1–)1.2–1.5 mm slightly from those that would only recognize (DNA
wide P. hartwegii based) phylogenetic characters. This approach
9b. Leaves slender, rigid or pliant, 0.7–1.1 mm makes it somewhat easier to use morphological
wide P. durangensis characters in this key to the amended subsections of
10a. Bark on trunk reddish brown to dark brown, the subgenus Strobus.
divided by very deep, longitudinal, black fis-
sures. Seed cones massive, 15–35 cm long. Seeds 1a. Fascicle sheaths of leaves persistent, not recoil-
8–10 mm long, with a 25–35 mm long wing ing; leaves connate, appearing as 1 but actually
P. devoniana with 3 (rarely 4) in a fascicle. Seed cones on rela-
10b. Bark grey-brown or brown, with plates or fis- tively very long, curved peduncles Subsect.
sures less deep relative to the trunk diam. Seed Nelsoniae: a single species P. nelsonii
cones smaller (maximum 20 cm long). Seeds 1b. Fascicle sheaths of leaves deciduous, scales first
4–7 mm long, with a 13–28 mm long wing 11 recoiling or not; leaves free (sometimes more or
11a. Seed cones solitary or in pairs, on distinct, less connate from the base but always distally
curved peduncles; seed scales thin and flex- free). Peduncles of seed cones much shorter
ible. Leaves very slender, pliant, 0.6–1(–1.1) mm than the cones 2
wide P. maximinoi 2a. Leaves in fascicles of only 5, these persisting
11b. Seed cones solitary or more often in whorls of on the branches for more than 10 and up to
2–6, on short peduncles or appearing sessile; 30 years, forming long ‘foxtail’ tufts; fascicle
seed scales thin or thick but rigid. Leaves rigid sheaths not recoiling and already absent in sec-
or pliant, 0.8–1.3 mm wide 12 ond year fascicles Subsect. Balfourianae
 2b. Leaves in fascicles of 1–5 (very rarely 6 or 7),
Key to the species of Pinus Subsection
these persisting 2–5 years (rarely as long as
Cembroides
8 years), with recoiling or early deciduous fas-
cicle sheaths 3 1a. Usually decumbent shrubs, multi-stemmed.
3a. Leaves flat, 1.5–5 mm wide, in fascicles of only Leaves predominantly in fascicles of 5 (very
2. Seeds with a long wing attached Subsect. rarely of 4 or 6), persisting 2–3 years. Seeds 5–7
Krempfiae: a single species P. krempfii mm long P. culminicola
3b. Leaves not flat, 0.5–2(–2.5) mm wide, in fasci- 1b. (Small) trees with at least a short single trunk
cles of 1–5 (very rarely 6 or 7); if with 2 leaves before branching. Leaves in fascicles of 1–5
then seeds without a wing attached 4 (rarely 6), persisting (3–)4–8 years. Seeds 10–18
616 4a. Bark on trunk (and branches) remaining mm long 2
smooth even in large trees, exfoliating with thin 2a. Leaves in fascicles of only 1 (rarely 2), 1.2–2.2(–
flakes creating a multi-coloured pattern some- 2.5) mm diam., terete unless with 2 in a fascicle;
what similar to bark of planes (Platanus) fascicle sheaths not recoiling P. monophylla
Subsect. Gerardianae 2b. Leaves in fascicles of (1–)2–5(rarely 6), (0.6–)
4b. Bark on trunk becoming rough and scaly at 0.7–1.5(–1.7) mm wide, not terete unless with 1
least in large trees, exfoliating irregularly, form- in a fascicle; fascicle sheaths recoiling or not 3
ing multi-layered small or large plates, or be- 3a. Scales of fascicle sheaths recoiling before fall-
coming fissured 5 ing. Integument of seeds thick and hard 4
5a. Umbos of seed cone scales terminal (at apex of 3b. Scales of fascicle sheaths deciduous without re-
scale). Leaves in fascicles of 5 (rarely 6 or 7) coiling. Integument of seeds thin, crushed eas-
Subsect. Strobi ily 5
5b. Umbos of seed scales dorsal (above apex of 4a. Leaves 2–4 cm long, predominantly in fascicles
scale). Leaves in fascicles of 1–5 (rarely 6) 6 of 2, sometimes in fascicles of 1 or 3 P. edulis
6a. Leaves 5–12(–14) cm long. Seed cones 5–25(–27) 4b. Leaves (2–)3–6(–8) cm long, predominantly in
cm long, if shorter than 8 cm only partly open- fascicles of 3, but commonly of 2 or 4 (rarely of
ing with thick but flexible seed scales 5) P. cembroides
Subsect. Rzedowskianae 5a. Leaves in fascicles of 2, sometimes in fascicles
6b. Leaves (1.5–)2–6(–8) cm long. Seed cones (2–) of 3, 0.8–1.1 mm wide. Seed cones (2–)2.5–4 cm
2.5–6(–7.5) cm long, always opening widely long P. remota
with thin, flexible seed scales 5b. Leaves in fascicles of (3–)4(–5), rarely in fasci-
Subsect. Cembroides cles of 2 or 6, (0.8–)1–1.5(–1.7) mm wide. Seed
cones 4–6 cm long P. quadrifolia
Key to the species of Pinus Subsection
Balfourianae Key to the species of Pinus Subsection
Gerardianae
1a. Leaves with small resin droplets drying as
white specks. Umbos on seed cone scales with a 1a. Leaves in fascicles of 4–5; 9–17 cm long; buds
strong, 4–10 mm long prickle P. aristata resinous. Seed cones on 1.5–2 cm long pedun-
1b. Leaves without small resin droplets. Umbos cles; seeds with a ca. 15 mm long wing
with a weak, 1–6 mm long prickle or unarmed P. squamata
2 1b. Leaves in fascicles of only 3; 5–10 cm long; buds
2a. Bark on the trunk breaking into small, irregular not resinous. Seed cones nearly sessile; seeds
plates, creating a tessellate pattern, cinnamon without a wing or wing rudimentary 2
or orange-brown. Seeds 8–10 mm long 2a. Seed cones 5–7 cm long; umbos of seed scales
P. balfouriana with a rigid prickle or spine; seeds 8–10 mm
2b. Bark on the trunk breaking into irregular, scaly long. Leaves up to 2 mm wide P. bungeana
ridges, shallowly to deeply fissured, reddish 2b. Seed cones 12–20(–23) cm long; umbos of seed
brown. Seeds 5–8 mm long P. longaeva scales unarmed; seeds 20–25 mm long. Leaves
1 mm wide P. gerardiana
 thick and rigid; seeds without a wing or wing
Key to the species of Pinus Subsection
rudimentary 6
Rzedowskianae
5b. Seed cones opening soon and fully, deciduous;
1a. Leaves in fascicles of 3, rarely of 4, rigid, 0.8–1.2 seed scales thin or thick; seeds with a wing at
mm wide; scales of fascicle sheaths early decid- least half the length of the seed, usually much
uous, not recoiling P. pinceana longer 9
1b. Leaves in fascicles of (3–)4–5, lax, 0.5–0.8 mm 6a. Leaves persisting on the branches (3–)5–6
wide; scales of fascicle sheaths recoiling before years. Pollen cones 6–10 mm long P. flexilis
dropping off 2 6b. Leaves persisting on the branches 2–3 years.
2a. Bark on the trunk thick, deeply fissured, dark Pollen cones at least 15 mm long 7
brown. Pollen cones small, 5 × 3 mm, purplish 7a. Seed cones 5–7 cm long, ovoid to ovoid-elliptic 617
before anthesis. Seed cones 10–15 cm long, with when closed. Leaves (3–)5–8(–9) cm long, 0.8–1
thin scales; seeds with a long wing attached mm wide P. amamiana
P. rzedowskii 7b. Seed cones 8–14 cm long, ovoid-oblong or con-
2b. Bark on the trunk breaking into square plates, ical-cylindric when closed. Leaves (5–)6–14 cm
not fissured, outer bark grey. Pollen cones 8–10 long, 1–1.5 mm wide 8
mm long, yellowish before anthesis. Seed cones 8a. Margins of seed scales recurved. Pollen cones
massive, (15–)17–25(–27) cm long, with very yellow. Young shoots occasionally pubescent
thick scales; seeds without a wing attached P. koraiensis
P. maximartinezii 8b. Margins of seed scales not recurved. Pol-
len cones greenish white with a reddish apex.
Young shoots always glabrous P. armandii
Key to the species of Pinus Subsection Strobi
9a. Leaves drooping or pendulous, straight but of-
The foliage characters in this subsection of pines ten with a sharp bend near the base. Bark on
are very similar and, however desirable in the situ- the trunk of large trees fissured longitudinally
ation where there are no cones, as in young trees, 10
they cannot be used reliably as first characters in the 9b. Leaves lax, pliant or rigid, spreading, straight
leads in most cases. Seed cones and sometimes pol- or (slightly) curved. Bark on the trunk of large
len cones are necessary and even these can be very trees not fissured longitudinally 11
similar; see the note at the end of this key. 10a. Young shoots glandular pubescent; leaves soon
deciduous, remaining only 1.5–2 years, pendu-
1a. Seed cones nearly sessile, persistent, serotinous. lous, 15–24 cm long P. bhutanica
Pollen cones reddish or red-purple 2 10b. Young shoots glabrous; leaves remaining on the
1b. Seed cones pedunculate, deciduous or some- shoots 2–3 years, drooping or sometimes pen-
times semi-persistent, opening soon or more dulous, (6–)10–18(–20) cm long P. wallichiana
slowly. Pollen cones yellow, greenish white or 11a. Seed cones in clusters of 4–8(–10), nearly ses-
sometimes tinged reddish at the apex 5 sile, persistent, ovoid or sub-globose when
2a. Shrubs, spreading with layering stems. Buds closed, 4–7 cm long P. parviflora
resinous. Seed cones 3–5 cm long P. pumila 11b. Seed cones solitary or in whorls of 2–4, pedun-
2b. Trees. Buds not resinous. Seed cones 5–10(–12) culate, deciduous, ovoid to long cylindrical,
cm long 3 variable but usually longer than 7 cm 12
3a. Leaves persisting on the branches for up to 12a. Seed cones with thick, rigid scales and these
8 years, 3–6.5 cm long P. albicaulis with thick, woody apophyses, large (but vari-
3b. Leaves persisting on the branches 2–4 years, able in one species) to 55–60 cm long 13
6–11(–13) cm long 4 12b. Seed cones with thin, flexible or more rigid
4a. Seed cones broadly ovoid, 5–8 cm long scales and these with thin or only partly thick-
P. cembra ened apophyses, (3–)5–40(–50) cm long 14
4b. Seed cones ovoid-conical, (5–)7–10(–12) cm 13a. Seed cones when mature at least 30 cm long,
long P. sibirica with 5–12 cm long peduncles. Bark on large
5a. Seed cones opening slowly and only slightly, trunks breaking into large plates
late deciduous to semi-persistent; seed scales P. lambertiana
 13b. Seed cones when mature 12–30(–60) cm long, the south of its current range and extended probably
on 1.5–2.5 cm long peduncles. Bark on large into Mexico.
trunks breaking into small plates P. strobiformis
14a. Seed scales near base (but well above the pe-
duncle) upturned or reflexed 15 Pinus albicaulis Engelm., Trans. Acad. Sci. St. Louis
14b. Seed scales near base (but well above the pe- 2: 209. 1863. Type: USA: Colorado, J. S. Newberry
duncle) incurved or more or less straight and s.n. (US No. 61140) (holotype US). Fig. 205
flat 17
15a. Seed cones 10–25(–30) cm long; seeds with a
Etymology
20–25 mm long wing. P. monticola
618 15b. Seed cones (3–)4–15(–17) cm long; seeds with a The Latin epithet albicaulis means ‘with a white
rudimentary or short wing to 16 mm long 16 stem’, referring to the light grey bark.
16a. Leaves 2.5–6 cm long; resin ducts in leaves me-
dial P. wangii
Vernacular names
16b. Leaves 4–18 cm long; resin ducts in leaves ex-
ternal, sometimes a few medial P. fenzeliana Whitebark pine, Scrub pine
17a. Seed cones with short, curved or obliquely
inserted peduncles, or more or less curved at
Description
base. Leaves remaining on the shoots 3–4 years
18 Trees to 10–15(–21) m tall, d.b.h. to 1–1.5 m. Trunk
17b. Seed cones with up to 4 cm long, straight pe- monopodial or with 2-several stems, straight or con-
duncles and more or less straight at base. Leaves torted, branching very low and much reduced in size
remaining on the shoots 2–3 years 19 at tree line. Bark thin, rough and scaly, breaking into
18a. Pollen cones 15–25 mm long. Seed cones 6–11 small, scaly plates, yellowish brown, weathering pale
cm long. Leaves 4–9 cm long, 0.6–1 mm wide grey, on young trees and branches smooth and grey-
P. morrisonicola ish white. Branches of first order contorted, usually
18b. Pollen cones 10–15 mm long. Seed cones (7–) short, ascending or spreading, persistent, of higher
8–15(–20) cm long. Leaves 7–10(–12) cm long, orders flexible, assurgent. Shoots often puberulent,
0.5–0.7 mm wide P. peuce with short, non-decurrent pulvini, pale red-brown,
19a. Seed cones (10–)15–40(–50) cm long, 7–15 cm ageing to pale grey-brown. Cataphylls 5–7 mm long,
wide when open. Leaves (8–)10–15(–18) cm subulate, scarious, brown, with entire margins, soon
long P. ayacahuite deciduous. Vegetative buds ovoid and acute; termi-
19b. Seed cones (5–)6–20(–25) cm long, 4–9 cm nal bud 8–10 mm long; lateral buds smaller, with
wide when open. Leaves (3–)5–12(–14) cm imbricate, appressed, subulate, red-brown scales.
long 20 Fascicle sheaths 8–12 mm long, with 6–7 loosely im-
20a* Buds slightly resinous [American species] bricate, orange-brown scales, deciduous and absent
P. strobus in second year’s fascicles. Leaves in fascicles of 5, in
20b* Buds not resinous [Vietnamese species] dense tufts towards ends of branchlets, persisting up
P. dalatensis to 8 years, mostly connivent, curved forward against
shoot, 3–6.5 cm long, 1–1.5 mm wide, with serrulate
* Both leaf lengths and seed cone lengths are ex- margins distally, acute, yellow-green on abaxial face,
tremely variable in these two species, ranging from white stomatal lines on both adaxial faces. Stomata:
(3–)5–12(–14) cm in the leaves and (5–)6–23(–25) cm leaves epistomatic or weakly amphistomatic, with
in the cones. These species, with disjunct ranges in 0–1 lines of stomata in grooves on abaxial face and
eastern North America, southern Mexico and parts 2–3 (intermittent) lines on each adaxial face. Pollen
of Guatemala (P. strobus), and Vietnam (P. dalaten­ cones crowded at the proximal end of a new shoot,
sis), are remarkably similar and probably closely re- spreading, subtended by light brown bracts, ovoid-
lated. The Mexican-Guatemalan populations are by oblong, 10–15 mm long, red, maturing to brown.
some accepted as a third species, Pinus chiapensis, Microsporophylls peltate, the distal part cordate to
but are here maintained as a variety of P. strobus, rounded, smooth, ca. 1 mm wide. Seed cones sub-
which during the Ice Ages occupied refugia far to terminal, solitary, in pairs or in whorls of 3, on very
short peduncles, persistent. Immature cones erect,
Conservation
ovoid, dark grey or purplish, maturing in two sea-
sons. Mature cones broadly ovoid to subglobose, 5–8 The most severe threat to this species at present is
× 4–7 cm. Seed scales ca. 50, remaining closed, con- infestation by the introduced rust Cronartium ribi­
cavo-convex, with seed cavities near base on adaxial cola, which has led to reductions of 80-90% in some
side, dark grey or purple, maturing to light brown. northern areas of its range, where there is a wetter
Apophysis thick, triangular to rhombic, transversely climate. Additional threats are predation by the bee-
keeled, straight or recurved but not reflexed at mid- tle Dendrococtonus ponderosae (attacking especially
cone, light brown, resinous. Umbo terminal, trans- weakened pines infested by the rust) and fire sup-
verse-triangular, acute, 5–6 mm wide, grey-brown, pression in National Parks leading to a successional
very resinous. Seeds obovoid, 7–11 × 4–7 mm, chest- change in vegetation. Attempts are being made to 619
nut brown. Seed wings absent when the seed is free select rust-resistant genetic variants and to adjust
from the scale. the fire regimes in National Parks to a more natural
frequency. This species has a very wide distribution
and not all populations are similarly effected.
Distribution
IUCN: VU [A1c]
W North America: Rocky Mountains, Cascade
Range, Sierra Nevada.
Uses
TDWG codes: 71 ABT BRC 73 IDA MNT ORE WAS
WYO 76 CAL NEV Whitebark pine’s main value is providing habitat
for wildlife. The wood is used locally to build log
cabins and for firewood. The edible seeds are diffi-
Ecology
cult to harvest due to the serotinous and resinous
Pinus albicaulis is a high altitude pine growing in cones; they were used by Native Americans only as
subalpine conifer forests up to the timberline. It oc- an emergency food source. Its aesthetic value is best
curs at altitudes up to 1350 m a.s.l. in the north and appreciated in situ in the high mountains; the hor-
3650 m a.s.l. in the south of its range. It is a very ticultural merits of this pine are low because of ex-
slow growing and long-lived tree, with often con- tremely slow growth and vulnerability to pathogens
torted stems and branches on exposed mountain such as white pine blister rust (Cronartium ribicola,
ridges. Whitebark pine occurs in a cold, moist, and Basidiomycota) and ‘late frost’ in parks and gardens.
snowy climatic zone exposed to high winds, where It is therefore scarcely known in cultivation.
few trees can compete. Most commonly it is associ-
ated with Abies lasiocarpa, Tsuga mertensiana, and
in regions with less abundant precipitation, Pinus Pinus amamiana Koidz., Bot. Mag. (Tokyo) 38:
contorta. Rain only occurs in the short summer 113. 1924. Pinus armandii Franch. var. amamiana
period (June-September), during the remainder of (Koidz.) Hatus., Mem. Fac. Agric. Kugos Univ. 1: 37.
the year sleet and snow prevail, and the snow pack 1974. Type not designated.
can cover small trees entirely except on windswept
ridges. The strong, flexible branches of P. albicaulis
Etymology
can withstand winds of hurricane force. Pinus al­
bicaulis co-evolved with a mountain bird, Clark’s This species was named after the Amami people,
Nutcracker (Nucifraga columbiana, Corvidae; see who inhabit the Ryukyu Islands.
also under P. cembra) in a mutualism concerning
seed dispersal. The bird removes the seeds from the Vernacular names
closed cones without damaging them and puts them
Amami pine; Amami-goyo, Yaku-tane-goyo (Japa-
away for later use in caches. Those that are forgot-
nese)
ten may germinate, often two to five trees thus grow
close together. No germination occurs without help
Description
from the bird and the bird even feeds its young with
the pine seeds. Red squirrels play a similar but lesser Trees to 20(–30) m tall; trunk to 2 m d.b.h. Bark
role in seed planting. smooth in young trees, greyish brown, becoming
 fissured and breaking into large and flaking plates
Distribution
on lower trunk, grey or blackish grey. Branches in
pseudo-whorls in young trees, forming a conical Japan, Kyushu (Yakushima, Tanegashima).
crown; in older trees crown widening and becoming TDWG codes: 38 JAP-KY
rounded and open. Foliage branches smooth, young
shoots puberulent, soon glabrous, grey-green, turn-
Ecology
ing grey-brown. Buds cylindrical, slightly resinous;
terminal bud 10–15 mm long; lateral buds smaller; Pinus amamiana occurs in exposed, open stands in
cataphylls ovate-linear, reddish brown. Leaves in re- often sparsely vegetated localities on rocky slopes at
mote fascicles of 5, held by a soon deciduous sheath between 100 m and 300 m a.s.l. on two smaller is-
620 of flimsy scales, persisting 2–3 years, slender and lands in the south of Japan.
spreading, (3–)5–8(–9) cm long, straight or slightly
curved, triangular in cross-section, 0.8–1 mm wide;
Conservation
stomata in lines on the two adaxial faces only. Pol-
len cones in long clusters, spirally arranged at base This rare species has an area of occupancy (AOO) of
of new shoots, cylindrical or ovoid-ellipsoid, 1.5–2.5 less than 100 km2; the total population size amounts
(–3) cm long, slender or stout, greenish white with a to fewer than 1500 trees (ca. 1000 on Yakushima)
slightly reddish apex. Seed cones solitary or in pairs, and is declining. These trees were formerly exploited
initially erect and green, becoming more or less for timber and regeneration is slow due to exposed
pendulous on stout, curved, 1–2 cm long peduncles, conditions. Nematodes accidentally introduced
opening only slightly, ovoid to ovoid-elliptic, 5–7 from the USA have caused increased mortality
cm long, 3–4 cm wide when opened, often resinous, among seedlings and saplings according to informa-
turning dark purplish brown. Seed scales woody, tion obtained by Tetsukazu Yahara of the Japanese
rigid, 1.5–2 cm long, 2–3 cm wide, curved slightly Plant Specialist Group (IUCN-SSC) in 1999.
inward; basal scales scarcely or not recurved, on IUCN: EN [C1]
the adaxial side with two marked depressions hold-
ing the seeds. Apophysis rhombic or more or less
Uses
rounded distally, thick woody, with a straight or
slightly recurved edge terminating in an inconspic- No recent uses have been recorded of this species;
uous, obtuse umbo, light yellowish brown or darker in the past its timber was exploited and used locally
red-brown. Seeds ellipsoid, 10–12 mm long, 5–6 mm for construction and carpentry. It is reported to be
wide, ca. 4 mm thick, grey to nearly black, wingless very rare in cultivation, but since it has frequently
or with a small ridge on abaxial margin, rarely with been referred to as P. armandii var. amamiana or
a rudimentary wing. even equated with that species (as it was considered
by E. H. Wilson, 1916), there may be trees in collec-
tions (arboreta etc.) that are misidentified. It may be
Taxonomic notes
somewhat more common in Japanese gardens.
The relationship of this species with the very similar
species P. armandii, P. fenzeliana and P. morrisonicola
Pinus aristata Engelm., Amer. J. Sci. Arts, ser. 2,
and the infraspecific taxa in this group of East Asian
34: 331. 1862. Pinus balfouriana Balf. var. aristata
pines belonging to subsection Strobus is in need
(Engelm.) Engelm., in Rothrock, Rep. US Geogr.
of further investigation using DNA sequence data.
Surv., Wheeler, 6, Bot.: 375. 1878; Pinus balfouriana
Pinus amamiana may be most distinct in its cones
Balf. subsp. aristata (Engelm.) Engelm., Trans. St.
and seeds, which seem to be more morphologically
Louis Acad. Sci. 4: 176. 1880. Type: USA: Colorado,
adapted to seed dispersal by birds, with small cones
C. C. Parry s.n. (lectotype MO). Fig. 206
and relatively large, virtually wingless seeds. How-
ever, as we have learned from DNA analysis of other
Etymology
pines with such cones (e.g. P. albicaulis and P. cembra)
strong selective pressure has determined the evolu- The species epithet is from Latin arista = ear (of
tion of these adaptations and they are not necessarily corn); it refers to the long, slender prickles or bris-
good indicators of phylogenetic relationships. tles on the cone scales causing the resemblance.
 TDWG codes: 73 COL 76 ARI 77 NWM
Vernacular names
Rocky Mountain bristlecone pine, Colorado bristle­
Ecology
cone pine, Hickory pine
Pinus aristata occurs in the subalpine and alpine
zones of the central Rocky Mountains; its altitudinal
Description
range is between 2500 m and 3400 m a.s.l. It is often
Trees, small to medium, height to 10–15 m, d.b.h. to found on extremely exposed slopes and ridges that
1–1.2 m. Trunk monopodial, straight or contorted, can be battered by severe snow and ice storms. It
often strongly tapering in old trees, branching very may occur in pure, scattered stands or together with
low and much reduced in size at tree line. Bark thin, Pinus albicaulis or Picea engelmannii. It is extreme- 621
shallowly fissured, breaking into flat, irregular ridg- ly slow growing and, although no trees have been
es, reddish brown, weathering grey. Branches of first found that quite match the maximum ages observed
order contorted, usually short, ascending or spread- in P. longaeva to the west, some individual trees are
ing, persistent, of higher orders flexible. Shoots pu- estimated to be in excess of 3000 years old.
berulent, with short, non-decurrent pulvini, pale
red-brown, ageing to grey. Cataphylls 5–7 mm long,
Conservation
subulate, scarious, brown, with entire margins,
soon deciduous. Vegetative buds ovoid and acute, IUCN: NT
resinous; terminal bud 8–10 mm long, lateral buds
smaller; with imbricate, appressed, subulate, red-
Uses
brown scales. Fascicle sheaths deciduous and absent
in second year’s fascicles. Leaves in fascicles of 5, in Rocky Mountain bristlecone pine is not a timber
dense tufts persisting up to 17 years, mostly conniv- tree due to unfit shape and extremely slow growth. It
ent, curved forward against shoot, 3–4.5 cm long, is in cultivation and grown as an ornamental in rock
0.8–1 mm wide, with white resin droplets that dry gardens and other suitable settings, especially in the
into small white specks or scales; margins entire or USA, and is commonly present in arboreta of the
distally serrulate; apex acute or subulate, leaf colour NE and NW USA, Canada and northern Europe.
glaucous green on the deeply grooved abaxial face, Although tolerant of poor, rocky soil and dry condi-
many white stomatal lines on both adaxial faces. tions, this species is often difficult to keep alive for
Pollen cones crowded at the proximal end of a new long due to susceptibility to pathogens under moist
shoot, spreading, subtended by light brown bracts, air conditions. As a species of high alpine habitats
ellipsoid or ovoid, ca. 10 mm long, mostly yellow in mid-continental climate, once warm weather sets
(a few red or bluish purple), maturing to brown. in it will flush and not expect frost again, as often
Microsporophylls peltate, the distal part cordate to occurs in more maritime climates of the north-
rounded, smooth, ca. 1 mm wide. Seed cones subter- ern hemisphere. A few cultivars have been named,
minal, solitary or in pairs, nearly sessile, falling after mostlly based on compact growth forms.
seed dispersal. Immature cones erect, ovoid, dark
purple, maturing in two seasons. Mature cones nar-
rowly ovoid becoming ovoid-cylindric when open, Pinus arizonica Engelm., in Rothrock, Rep. US
6–11 × 4–6 cm. Apophysis on seed scales thick, tri- Geogr. Surv., Wheeler, 6, Bot.: 260. 1879.
angular to rhombic, transversely keeled, recurved,
brown. Umbo dorsal, transverse-triangular at base,
Etymology
terminating in a slender, brittle, recurved and 4–10
mm long prickle. Seeds obliquely obovoid, 5–6 × The species epithet refers to Arizona, from where it
3–4 mm, grey-brown to near black. Seed wings 10– was first described.
14 mm long.
Vernacular names
Distribution
Arizona pine
USA: N Arizona, Colorado, N New Mexico.
622

Pl ate 2 6 . Pinus arizonica var. arizonica. 1. Habit of tree. 2. Leaves. 3. Seed cones, one green, one ripe.
 S Durango, scattered in Coahuila, NE Zacatecas and
Description
Nuevo León.
Trees to 30–35 m tall, d.b.h. to 1–1.2 m. Trunk mo- TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CO
nopodial, erect, straight. Bark thick, rough, scaly, MXE-CU MXE-DU MXE-NL MXE-ZA MXN-SI MXN-
breaking into large plates divided by broad, shal- SO
low or deep fissures, reddish brown to dark brown;
outer bark grey. Branches spreading or ascending,
Ecology
lower branches subpendulous, slender. Crown in
old trees rounded or flat-topped, open or dense. Pinus arizonica is forming pure stands or is more
Shoots at first orange-brown or glaucous, rough commonly mixed with Quercus spp., other pines,
with persistent, decurrent pulvini. Cataphylls up to e.g. Pinus engelmannii, P. strobiformis, and occa- 623
15 × 3–4 mm, subulate, scarious, soon reflexed, with sionally Juniperus flaccida at lower or J. deppeana
erose-ciliate margins and caudate apex, dark brown. at higher altitudes. It grows on various substrates,
Vegetative buds ovoid to ovoid-acute; terminal bud but the best stands are in valleys and on mesas with
15–30 × 10–15 mm; lateral buds smaller, not resin- deep soil, in moderately dry to mesic forest with
ous, brown; the scales spreading or recurved at apex, light winter frost occurring, at altitudes from (1300–
with long-ciliate, white or brown margins. Fascicle )2000–2700(–3000) m a.s.l. Annual precipitation is
sheaths initially long, 15–30 mm, reduced to ca. 10 low to moderate, from 700 mm to 900 mm, mostly
mm. Leaves in fascicles of 3–5, persisting 2–3 years, falling during the winter months.
rigid or pliant, straight or slightly curved and twisted,
(8–)10–20(–25) cm long, 0.9–1.8 mm wide; margins
Conservation
serrulate; apex acute to pungent; leaf colour light yel-
lowish green or glaucous green. Stomata on all faces Pinus arizonica is an important constituent of the
of leaves, in (3–)4–8(–12) lines on the convex abax- pine and pine-oak forests of northern Mexico, es-
ial face and in (3–)4(–5) lines on each adaxial face. pecially in the Sierra Madre Occidental. As such it
Pollen cones ovoid-oblong to cylindrical, 1.5–2 cm is heavily exploited as a timber tree in much of its
× 5 mm, yellowish to yellowish brown. Seed cones range, especially in the more accessible regions, and
solitary, in pairs or whorls of 3–5, on short, curved, stands with large trees in these localities are now
persistent peduncles retaining a few scales on the rare (Perry, 1991). However, there is no evidence that
branches. Mature cones seemingly sessile, ovoid either its range, nor the number of trees surviving,
or broadly ovoid with parted scales, asymmetrical have declined and so under IUCN criteria this spe-
at obliquely flattened base, often slightly curved, cies is not threatened with extinction.
(4.5–)5–10(–14) × 3.5–6(–8) cm when open. Seed
scales thick, rigid, broadly oblong, nearly straight
Uses
or recurved especially near base, dark purplish to
blackish brown on abaxial side, light brown with In Mexico, Pinus arizonica is a major timber tree,
faint marks of seed wings on adaxial side. Apophysis often compared (or equated) with P. ponderosa in
rhombic to pentagonal in outline, mostly symmet- the USA. The uses of its wood are similar. Timber
rical around cone, transversely keeled, ochraceous of large sizes and high grade is sawn for light con-
to light reddish brown. Umbo dorsal, raised and struction like window frames, doors, stairs, flooring,
slightly recurved or nearly flat, rhombic, transverse- sidings, panelling, and veneers and for furniture,
ly keeled, 4–8 mm wide, grey, with a minute, decidu- cabinetwork, boxes, and woodware. There are many
ous prickle. Seeds obliquely ovoid, slightly flattened, local or regional sawmills where the logs are sawn
4–7 × 3–5 mm, light brown with black spots; wings and relatively little is exported. The resource has
obliquely ovate, 12–20 × 4–8 mm, semi-transparent, become much reduced by overexploitation and it
greyish brown with black tinge. is difficult to implement more sustainable forestry
methods in the short run that would reverse the
downward trend. This pine is not known to be in cul-
Distribution
tivation outside a few trial plantations, but perhaps
SW USA: Arizona, New Mexico; in Mexico main- an occasional tree from southern Arizona identified
ly in the Sierra Madre Occidental, southwards to
 as P. ponderosa planted in parks or gardens belongs on the convex abaxial face and in (3–)4 lines on each
to this species. adaxial face (abaxial number of lines correlated with
width of leaf). Seed cones ovoid to broadly ovoid, of-
3 varieties are recognized: ten slightly curved, 5–10(–12) × 4–6 cm when open.
Seeds obliquely ovoid, slightly flattened, 5–7 × 4–5
Pinus arizonica Engelm. var. arizonica. Pinus mm. Seed wings obliquely ovate, 15–20 × 6–8 mm.
ponderosa Douglas ex C. Lawson var. arizonica
(Engelm.) Shaw, [Pines Mexico] Publ. Arnold
Distribution
Arbor. 1: 24. 1909; Pinus ponderosa Douglas ex
C. Lawson subsp. arizonica (Engelm.) E. Murray, Mexico: mainly in Durango, but scattered north-
624 Kalmia 12: 23. 1982. Type: USA: Arizona, Pima Co., ward in the Sierra Madre Occidental into Chihua-
Santa Rita Mts., J. T. Rothrock 652 (holotype MO). hua and Sonora.
Pl. 26 TDWG codes: 79 MXE-CO MXE-CU MXE-DU MXN-
SO
Description
Conservation
Trees, usually tall, height to 30–35 m. Leaves in fas-
cicles of 3–5 (predominantly 3–4), rigid to slightly IUCN: LC
lax, straight or slightly curved and twisted, (8–)10–
20(–23) cm long, 0.9–1.4(–1.6) mm wide. Stomata Pinus arizonica Engelm. var. stormiae Martínez,
on all faces of leaves, in (3–)4–8 lines on the convex Pinos Mexic., ed. 2: 295. 1948. Pinus ponderosa
abaxial face and in (3–)4(–5) lines on each adaxial Douglas ex C. Lawson var. stormiae (Martínez)
face (abaxial number of lines correlated with width Silba, Phytologia 68: 59. 1990. Type: Mexico:
of leaf). Seed cones ovoid to broadly ovoid, often Coahuila, General Cepeda, Sierra la Cocordia,
slightly curved, (4.5–)5–7 × 3.5–6 cm when open. M. Martínez 3455 (lectotype MEXU).
Seeds obliquely ovoid, slightly flattened, 4–6 × 3–3.5
mm. Seed wings obliquely ovate, 12–15 × 4–6 mm.
Description
Trees of medium size, height to 15–20 m. Leaves
Distribution
in fascicles of 3–4(–5), thick, coarse, rigid, usually
SW USA (Arizona, New Mexico); mainly in the Si- curved and twisted, 14–25 cm long, 1.4–1.8 mm
erra Madre Occidental of Mexico southwards to S wide. Stomata on all faces of leaves, in 8–12 lines on
Durango, scattered in Coahuila, NE Zacatecas and the convex abaxial face and in (3–)4 lines on each
Nuevo León. adaxial face (abaxial number of lines correlated with
TDWG codes: 76 ARI 77 NWM 79 MXE-CO MXE- width of leaf). Seed cones ovoid to ovoid-oblong,
CU MXE-DU MXE-NL MXE-ZA MXN-SI MXN-SO often curved, (4.5–)5–10(–14) × 3.5–6(–8) cm when
open. Seeds obliquely ovoid, slightly flattened, 5–6 ×
3.5–4 mm. Seed wings obliquely ovate, 12–16 × 6–8
Conservation
mm.
IUCN: LC
Distribution
Pinus arizonica Engelm. var. cooperi (C. E. Blanco)
Farjon, Flora Neotropica 75: 107. 1997. Pinus cooperi Mexico: mainly in S Nuevo León, a few localities in
C. E. Blanco, Anales Inst. Biol. Univ. Nac. México S Coahuila, Zacatecas and possibly San Luís Potosí.
20: 185. 1950. Type: Mexico: Durango, El Salto, (coll. TDWG codes: 79 MXE-CO MXE-NL MXE-SL MXE-
C. E. Blanco) M. Martínez 3442 (lectotype A). ZA

Trees, usually tall, height to 30–35 m. Leaves in fas-

Conservation
cicles of (3–)4–5, predominantly 5, rigid to slightly
lax, usually curved, (5–)6–10(–12) cm long, 1.0–1.3 IUCN: LC
mm wide. Stomata on all faces of leaves, in 4–7 lines
fig ure 17 7. Nothotsuga longi­
bracteata in Nan Ling Mts. Hunan,
China

figure 17 8. Nothotsuga longi­

bracteata seed cones (photo Y. Liu)

figure 179. Papuacedrus papuana var.

­papuana pollen cones (photo D. White)

figure 182. Parasitaxus usta foliage

and seed cones (photo W. Baker)

fig ure 184. Phyllocladus aspleniifolius

­foliage and pollen cones

fig ure 181. Parasitaxus usta in

New Caledonia (photo W. Baker)

figure 180. Papuacedrus papuana

leaves and seed cones (photo D. White)

figure 183. Pherosphaera hookeriana

in Tasmania, Australia
fig ure 185. Phyllocladus
­hypophyllus canopy on
Mt Kinabalu, Borneo

figure 187. Phyllocladus

­trichomanoides var. alpinus foliage

fig ure 186. Phyllocladus

­hypophyllus glaucous foliage on
Mt. Kinabalu 3100 m

figure 188. Picea chihuahuana

green and ripe seed cones

fig ure 190. Picea likiangensis

var. likiangensis seed cones (photo
P. de Spoelberch)

figure 189. Picea likiangensis

var. likiangensis young seed cones

fig ure 192. Picea morrisonicola

tree in Taroko N.P., Taiwan

figure 193. Picea morrisoni­

cola trunk with bark

figure 191. Picea likiangensis var.

likiangensis seed cones
 figure 195. Picea
orientalis seed cones
f i g u re 1 94 . Picea orientalis
foliage and pollen cones

f i g u re 1 9 6 . Picea schrenkiana subsp. tianschanica in Kirgyzstan

figure 197. Picea

f i gu re 1 9 8 . Picea sitchensis tree in ­schrenkiana subsp.
­Olympic N.P., Washington, USA tianschanica tree in Kirgyzstan

figure 199. Picea

smithiana pollen cones

figure 2 00. Picea

smithiana seed cone
from bud
 f i g u re 2 03. Pilgerodendron uviferum pollen cones

figure 2 04. Pilgero­

f i g u re 2 01 . Picea dendron uviferum
­smithiana seed cone foliage and seed cones

figure 2 02. Picea

wilsonii pollen cones

figure 2 05. Pinus

albicaulis foliage and
pollen cones

figure 2 07. Pinus armandii

var. armandii seed cone

figure 2 0 6. Pinus
aristata foliage

fig ure 2 09. Pinus ­ayacahuite var. veitchii

foliage and seed cones

figure 2 08. Pinus

attenuata seed cones
 figure 2 11.
Pinus bungeana
trunk with bark

figure 2 10.
Pinus balfouri­
ana in the ­Sierra
Nevada, Califor-
nia, USA

fig ure 2 12. Pinus

bungeana seed cone

fig ure 2 13. Pinus cembra

in the Alps, Switzerland

f i g u re 2 1 4 . Pinus cembra
foliage and seed cones

f i g u re 2 15 . ­Pinus
­cembroides var.
­cembroides in
Hidalgo, Mexico

f i g u re
2 16 . ­Pinus
cembroides var.
­cembroides bark
f i gu re 2 1 7. Pinus cembroides figure 2 19 . Pinus
subsp. orizabensis seed cones coulteri pollen cones

figure 2 18. ­Pinus

contorta var.
­murrayana in the
Sierra Nevada, Cali-
fornia, USA

figure 223. ­Pinus

durangensis in
­Durango, Mexico
(photo C. Hughes)
f i gu re 2 2 0 . Pinus coulteri figure 2 2 1. Pinus
foliage and seed cones ­culminicola foliage

f i gu re 2 22. Pinus figure 2 2 4 . Pinus

­devoniana foliage engelmannii foliage
 figure 22 8. Pinus
l­ongaeva on Telescope
Peak, Death Valley N.P.,
California

figure 22 5. Pinus hartwegii

on Pico de Orizaba, Veracruz,
Mexico

figure 22 6. Pinus
­heldreichii seed cones
figure 230. Pinus maximartinezii
seed cone and seedling

figure 231. Pinus monophylla

foliage and seed cones

figure 229. Pinus maximartinezii

in Zacatecas, Mexico

figure 227. Pinus latteri forest in

­Thailand (photo H. Hazebroek)
f i gu re 2 32. Pinus monophylla leaves

figure 233. Pinus muricata figure 23 4 . Pinus

pollen cones muricata seed cones

f i g u re 2 35. Pinus nelsonii seed cone

f i gu re 2 3 8 . Pinus patula var. patula

tree in Oaxaca, Mexico

figure 236. Pinus nigra subsp. laricio figure 239. Pinus

trunk with bark patula var. patula leaves

figure 237. Pinus nigra subsp. salzmannii

seed cones
Pinus armandii Franch., Nouv. Arch. Mus. Hist.
Distribution
Nat., sér. 2, 7: 95. 1884.
China, Anhui, Chongqing, S Gansu, Guizhou,
Guangxi, Hainan, Henan, Hubei, S Shaanxi, Sichuan,
Etymology
Yunnan, extreme SE Xizang [Tibet]; N Myan­mar
This species was named after the French missionary [Burma]; Taiwan.
and naturalist Armand David (1826–1900), an avid TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
explorer of China. SC CHC-YN CHH CHN-GS CHN-SA CHN-SX CHS-AH
CHS-HE CHT 38 TAI 41 MYA
Vernacular names
633
Ecology
Armand’s pine; hua shan song (Chinese)
Pinus armandii occurs in mountains at altitudes
from 900 m to 3500 m a.s.l.; usually in association
Description
with other conifers and seldom in pure stands. Com-
Trees to 30–35 m tall; trunk to 1 m d.b.h. Bark smooth mon conifer genera in these mixed forests are Abies,
in young trees, becoming fissured and breaking into Picea, Pseudotsuga, and in SW China also Larix.
large and more or less rectangular plates on the low- More often than these conifers, the pines tend to oc-
er trunk, grey or purplish grey. Branches in pseudo- cupy rocky areas with thin soils where other trees,
whorls in young trees, forming a conical crown, in among them angiosperms, are less competitive.
older trees crown widening and becoming rounded
and open. Foliage branches smooth, young shoots
Uses
glabrous, green, grey-green, or grey-brown, some-
times glaucous, turning brown. Buds cylindrical, This pine is a notable ornamental tree in China and
slightly resinous; terminal bud 10–15 mm long; later- it was introduced to France by Armand David in
al buds smaller. Leaves in remote fascicles of 5, some- 1895. It remains an uncommon tree in gardens and
times 6–7, held by a soon deciduous sheath of flimsy parks outside China, but has spread to arboreta in
scales, persisting 2–3 years, slender and spreading or many parts of the world. As a timber tree it is of lim-
drooping, (5–)8–14 cm long, often curved near base, ited value and exploited only for local use.
triangular in cross-section, 1–1.5 mm wide; stomata
in lines on the two adaxial faces only. Pollen cones 3 varieties are recognized:
in long clusters, spirally arranged at base of new
shoots, cylindrical or ovoid-ellipsoid, 1.5–2.5(–3) cm Pinus armandii Franch. var. armandii. Type not
long, slender or stout, greenish white with a slightly designated. Fig. 207
reddish apex. Seed cones solitary or in pairs, initially
erect and green, becoming more or less pendulous
Description
on stout, curved, 2–3 cm long peduncles, falling late
after seed dispersal, conical-cylindric, 8–14 cm long, New shoots green or grey-green. Pollen cones stout,
5–8 cm wide when opened, often resinous, turning 1.5–2 cm long, ovoid-ellipsoid. Apophyses in mature
variously brown. Seed scales woody, rigid, 3–4 × 2.5– cones thick woody, rhombic, not recurved or only
3 cm, curved slightly inward,basal scales scarcely or the umbo slightly recurved, yellowish brown or light
not recurved, on the adaxial side with two marked brown. Seeds dark brown to nearly black.
depressions holding the seeds. Apophysis rhombic
or triangular, often thick woody, with a straight or
Distribution
slightly recurved edge terminating in an inconspic-
uous, obtuse umbo, light yellowish brown or darker China: Chongqing, S Gansu, Guizhou, Hainan,
red-brown. Seeds obovoid, 10–15 mm long, 6–10 Henan, Hubei, Shaanxi, Sichuan, Yunnan, extreme
mm wide, light or dark brown to nearly black, wing- SE Xizang [Tibet]; N Myanmar [Burma].
less or with a small ridge on abaxial margin, rarely TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
with a rudimentary wing. SC CHC-YN CHH CHN-GS CHN-SA CHN-SX CHS-HE
CHT 41 MYA
 Pinus armandii Franch. var. mastersiana (Hayata)
Conservation
Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25 (19): 217, f.
IUCN: LC 8. 1908. Pinus mastersiana Hayata, Gard. Chron.,
ser. 3, 43: 194. 1908; Pinus armandii Franch. subsp.
Pinus armandii Franch. var. dabeshanensis mastersiana (Hayata) Businský, Acta Pruhoniciana
(W. C. Cheng & Y. W. Law) Silba, Phytologia 68: 47. 68: 13. 1999. Type: Taiwan: Nantou, Chia-i Pref.,
1990. Pinus dabeshanensis W. C. Cheng & Y. W. Law, Yu-Shan, [“Mt. Morrison”], G. Nakahara s.n., 3 Oct
Acta Phytotax. Sin. 13 (4): 85. 1975; Pinus fenzeliana 1905 (holotype TI).
Hand.-Mazz. var. dabeshanensis (W. C. Cheng &
Y. W. Law) L. K. Fu & Nan Li, Novon 7 (3): 262.
Description
634 1997. Type: China: Anhui, Yuexi, Laibangmen,
C. Q. Zhang 1 (holotype PE). New shoots grey-brown. Pollen cones when fully
grown slender, cylindrical, 2–3 cm long. Apophyses
usually slightly recurved towards apex, brown or
Description
red-brown when mature. Seed colour not recorded.
Leaves 5–14 cm long. Apophysis of seed scales thick
woody. Seeds light brown, with a rudimentary wing.
Distribution
Taiwan (Mt. Alishan, Mt. Yushan).
Taxonomic notes
TDWG codes: 38 TAI
In Flora of China 4: 24 (1999) this taxon is treated
as a variety of Pinus fenzeliana Hand.-Mazz. Farjon
Conservation
(1998, [2001], 2005b) has maintained it under P. ar­
mandii. Although the thicker seed scales and rudi- This variety of ‘soft pine’ is the only representative
mentary seed wings are most similar to P. armandii, of Pinus subsection Strobi native to Taiwan and the
we also know that these are adaptive traits that may timber of pines in this subsection is very much su-
evolve independently in species with zoochorous perior for high grade carpentry and furniture mak-
seed dispersal. Careful DNA analysis may be the ing. As a result, the resource has been overexploited
only solution to the problem of relationship of this and the population has dwindled. At present, several
taxon; it has so far not been included in a compre- remaining stands are protected within Yushan Na-
hensive analysis of Pinus section Strobus. tional Park.
IUCN: EN (A1c)
Distribution
Uses
China: Anhui (Jingzhai, Yuexi), Henan
(Shangcheng), Hubei (Luotian, Yingshan). An important timber tree, but a very limited re-
TDWG codes: 36 CHC-HU CHS-AH CHS-HE source.

Conservation
Pinus attenuata Lemmon, Mining Sci. Press 64: 45.
This variety is known from disjunct localities in a 1892. Type: USA: California, Berkeley, California,
limited area where three provinces meet and where J. G. Lemmon s.n. (lectotype UC). Fig. 208
deforestation is known to occur and has occurred
for a long time. In Flora of China it is called “an en-
Etymology
dangered plant” but it is not clear whether this is
based on IUCN criteria. The species epithet refers to the tapering (attenuate)
IUCN: VU (B1+2a-b) shape of the closed seed cone.

Vernacular names
Knobcone pine, Narrowcone pine
 Description Ecology
Trees to 15–20(–25) m tall, d.b.h. to 40–50 cm. This species is a fire successional tree with extremely
Trunk monopodial, straight or sometimes curved. persistent and serotinous seed cones on dry slopes
Bark thin, rough, scaly, breaking in small, rectangu- in chaparral or similar vegetation, also on rocky out-
lar plates, greyish brown to grey. Branches spread- crops with little other plant growth. In the northern
ing to assurging, forming an irregular or rounded, part of its range, where trees grow somewhat taller,
open crown. Shoots often multi-nodal, with short it is often mixed with several species of oak (Quercus
decurrent, persistent pulvini. Cataphylls subulate, spp.). In the USA its altitudinal range is from 300–
scarious, tapering into a caudate apex; margins 1200(–1700) m a.s.l. In Mexico it is found mainly
erose-ciliate, dark brown. Vegetative buds ovoid- from 250–600 m a.s.l. and close to the coast. 635
acute; terminal bud 15–30 × 10–15 mm; lateral buds
smaller, resinous, brown; the scales appressed, sub­
Conservation
ulate, scarious, with erose, hyaline margins. Fascicle
sheaths initially 10–18 mm long, reduced to 3–6 mm. IUCN: LC
Leaves in fascicles of 3, rarely 2, persisting 2–3 years,
rigid, straight, (8–)10–12(–14) cm long, 1.0–1.5 mm
Uses
wide; margins serrulate; apex acute; leaf colour light
green or slightly glaucous green. Stomata on all fac- Knobcone pine has little or no value as a timber tree.
es of leaves, in 8–12 lines on the convex abaxial face Its erratic growth habit and generally small size do
and in 3–5 lines on each adaxial face. Pollen cones not yield quality timber; its dry habitat ensures slow
ovoid-oblong to cylindrical, 1.5–2 cm × 5–7 mm, yel- growth to any useful size, although on establishment
lowish to yellowish brown. Seed cones subterminal, after fire its initial growth rate is quite rapid. Cross
solitary or more often in pairs or whorls of 3–5, on sections of stems with inclosed whorls of cones have
short, stout, recurved peduncles, very persistent on sometimes been polished and offered as curios in lo-
main stem and branches. Mature cones seemingly cal woodcraft shops. Its horticultural merits are like-
sessile, reflexed, extremely serotinous, ovoid-oblong wise not esteemed and it is therefore restricted to a
to ovoid-attenuate, slightly curved, asymmetrical, few arboreta and pineta, mainly in regions with a
8–15 × 3.5–6 cm when closed (up to 8 cm wide when suitable, Mediterranean type climate, or at least one
open). Seed scales thick, rigid, oblong, straight or with warm, dry summers and mild winters. In Cali-
slightly curved when parted, purplish to reddish fornia, hybridization experiments have resulted in
brown. Apophysis rhombic to pentagonal in out- hybrids with P. radiata (known as P. ×attenuradiata
line, transversely keeled, from slightly raised on Stockwell & Righter) and with P. muricata. All three
lower side of the cone to conical and curved on up- species belong to a group of pines informally known
perside, especially towards base of cone (there more as the ‘California closed-cone pines’ and are indeed
than 10 mm long), ochraceous or yellowish brown, closely related.
slightly lustrous, weathering grey. Umbo dorsal,
rhombic, from almost flat to pyramidal or unciform
in conjunction with development of the apophysis, Pinus ayacahuite Ehrenb. ex Schltdl., Linnaea 12:
curved towards cone apex, up to 5 × 5 mm, dark- 492. 1838.
er than apophysis; prickle absent. Seeds obliquely
ovoid, flattened; apex slightly acute, 5–7 × 3.5–4.5
Etymology
mm, blackish grey. Seed wings oblong, with one side
curved, 12–18 × 5–7 mm, yellowish to grey-brown. The species epithet takes up one of the indigenous
vernacular names for this pine.
Distribution
Vernacular names
Mexico: Baja California Norte; USA: California,
S. Oregon. Mexican white pine; acalote, ayacahuite, ocote gre-
TDWG codes: 73 ORE 76 CAL 79 MXN-BC tado, pinabete, pino gretado (Mexico, Guatemala)
 Description Ecology
Trees to 40–45(–50?) m tall, d.b.h. to 1.5–2 m. Pinus ayacahuite forms emergent trees in mixed
Trunk monopodial, straight. Bark rough, scaly, di- montane conifer forest on mesic sites, or grows in
vided into small, rectangular scales, exfoliating with groups or small groves; reaching its greatest extent
small flakes, grey-brown, weathering grey. Branches in Chiapas and W Guatemala. The best stands are
spreading horizontally, but lower branches some- on loamy, well drained soils. Its altitudinal range is
what drooping; crown pyramidal to broad coni- (1500–)1900–3200(–3600) m a.s.l.
cal. Shoots mostly glabrous or current year’s shoots
brownish puberulent, grooved; pulvini prominent,
Uses
636 short decurrent. Cataphylls small, 5–10 mm long,
subulate, with erose margins, dark brown, soon de- Pinus ayacahuite and its variety P. ayacahuite var.
ciduous. Vegetative buds small; terminal bud 8–15 × veitchii are both very important and highly sought
5–8 mm, ovoid-oblong; lateral buds smaller, ovoid, after timber trees as they are the tallest growing
not or only slightly resinous; the scales subulate- and most regular shaped representatives of the so-
linear, acute, free, dark purplish brown. Fascicle called soft pines in Mexico and Mesoamerica. The
sheaths 15–20(–25) mm long, deciduous. Leaves wood is used for construction, carpentry, furniture
in fascicles of 5, very rarely 6, persisting 2–3 years, and utilities such as containers, pallets and crates as
straight or slightly curved or twisted, not drooping, well as wood pulp. This Mexican pine with its large,
(8–)10–15(–18) cm long, 0.7–1.0 mm wide; margins pendent cones is uncommon in cultivation and re-
(weakly) serrulate; apex acute; leaf colour green or stricted to a few arboreta and private parks or gar-
grey-green with two whitish stomatal bands. Sto- dens with rare trees. Hybrids have been produced by
mata on the two adaxial faces only, in two dense bringing this species together with its close relatives
bands of 3–6 intermittent lines. Pollen cones ovoid in the subsection Strobi, e.g. P. flexilis, P. monticola,
to short cylindrical, 7–10 mm long, yellow, turn- P. strobus and P. wallichiana, the hybrid with the lat-
ing orange-brown. Seed cones towards end of main ter is known as P. ×holfordiana A. B. Jacks. and can
branches, subterminal, solitary or more often in also be found planted in some tree collections. For-
pairs or whorls of 3–4 on short (–25 mm) pedun- estry plantations with these trees outside their native
cles which usually fall with the cone. Mature cones ranges, e.g. in western Europe, have been unsuc-
pendulous, cylindrical, curved, tapering towards cessful due to the risk of infestation with White pine
apex, (10–)15–40(–50) cm × 7–15 cm when open. blister rust (Cronartium ribicola, ­Basidiomycota),
Seed scales spreading, thin, flexible or more rigid, which particularly affects introduced pine species
straight, the proximal scales recurved, transversely of this subsection. The fungus resides in alternative
nearly flat or concavo-convex, 5–7 cm long, rugose hosts; in this case species of Ribes, native shrubs in
or grooved, dull brown or dark reddish brown. much of Europe, which do not suffer damage. Eradi-
Apophysis irregularly trullate, recurved or reflexed cation of native flora in forestry is no longer accept-
towards apex, or apex curved up below umbo, lon- able practice in the EU and foresters have to look for
gitudinally grooved, dull light brown, very resinous. other tree species to grow.
Umbo terminal, broadly triangular, obtuse, 5–10
mm wide. Seeds in two cup-like depressions at base 2 varieties are recognized:
of scale, obovoid, laterally flattened, 8–15 × 6–9 mm,
dull brown with black spots. Seed wings oblique, Pinus ayacahuite Ehrenb. ex Schltdl. var. ayaca­
with a curved side or broad truncate, 20–35 × 8–12 huite. Type: Mexico: Hidalgo, Omitlan de Juarez,
mm, light brown with dark striations or dark brown. Montes del Lucero, near Hacienda de Guerrero,
C. A. Ehrenberg 647 (lectotype MO).
Distribution
Mexico: from Central Mexico southwards to Chia- Description
pas; Guatemala; Honduras; El Salvador.
Seed cones pendulous, cylindrical, curved, taper-
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
ing towards apex, (10–)15–40 cm long, 7–15 cm wide
PU MXC-TL MXE-GU MXE-HI MXE-QU MXG-VC
when opened. Seed scales ca. 100–120, thin, flexible;
MXS MXT-CI 80 ELS GUA HON
apophysis thin, trullate, usually recurved, rarely re- Pinus balfouriana Balf., in Murray, Bot. Exped.
flexed at apex. Seeds 8–10 × 6–8 mm. Seed wings Oregon 8: 1. 1853. Type: USA: California, Shasta
20–35 × 8–12 mm, tapering towards apex. Co., Mt. Shasta, [“On a range of mountains
between Chastey (Castey) and Scotts Valley (River)
lat. 41° 50”], J. Jeffrey 618 (holotype E). Fig. 210
Distribution
Central America: El Salvador, Guatemala, Hondu- Pinus balfouriana Balf. subsp. austrina Bruijn &
ras; S Mexico: northwards to Hidalgo. J. Mastrog., Syst. Bot. 5 (1): 102. 1980; Pinus balfou­
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC- riana Balf. var. austrina (Bruijn & J. Mastrog.) Silba,
PU MXC-TL MXE-HI MXG-VC MXS-GR MXS-MI Phytologia Mem. 7: 48. 1984.
MXS-OA MXT-CI 80 ELS GUA HON 637
Etymology
Conservation
This species was named after (but not by) John Hut-
IUCN: LC ton Balfour (1808–1884), who for bibliographic rea-
sons is to be credited with the authorship.
Pinus ayacahuite Ehrenb. ex Schltdl. var. veitchii
(Roezl) Shaw, [Pines Mexico] Publ. Arnold Ar-
Vernacular names
bor. 1: 10. 1909. Pinus veitchii Roezl, Cat. Grain.
Conif. Mexic.: 32. 1857. Pinus strobiformis Engelm. Foxtail pine
subsp. veitchii (Roezl) Frankis, Int. Dendrol. Soc.
Yearb. 2008. 2009. Type: Mexico: México, Valle de
Description
México, Volcán Popocatepetl, [“ridge between Mts.
Popocatepetl and Iztaccihuatl”], C. G. Pringle s.n. Trees to 22 m tall, d.b.h. to 2.5 m. Trunk monopodial,
(neotype A). Fig. 209 straight or slightly contorted, massive. Bark break-
ing into irregular, small plates forming tesselated
patterns, or becoming deeply fissured, cinnamon
Description
or orange-brown, weathering grey in the northern
Seed cones pendulous, broadly cylindrical, curved, population. Branches of first order short and con-
often very large, 15–50 cm long, 10–15 cm wide when torted, ascending or spreading, persistent; branches
opened. Seed scales ca. 100–150, not flexible; apo- of higher orders flexible and often pendent. Shoots
physis thick, trullate, with an extremely elongated, glabrous or puberulent, with short, non-decurrent
abruptly narrowed, reflexed distal part and some- pulvini, red-brown, ageing to yellowish grey or grey.
times an upturned apex. Seeds 10–15 × 7–9 mm. Cataphylls 5–7 mm long, subulate, scarious, brown,
Seed wings up to 2 × as long as the seed, oblique, with entire margins, soon deciduous. Vegetative
broad truncate or tapering to a rounded apex. buds ovoid and acute, resinous; terminal bud 8–10
mm long; lateral buds smaller; with imbricate, ap-
pressed, subulate, red-brown scales. Fascicle sheaths
Distribution
deciduous and absent in second year’s fascicles.
Mexico: in Guanajuato, Querétaro, Hidalgo, Ver- Leaves in fascicles of 5, in dense tufts persisting up to
acruz, México, Puebla, Tlaxcala, Morelos, and Mi- 30 years, mostly connivent and spreading outward
choacán. or curved forward along shoot, (1.5–)2–4 cm long,
TDWG codes: 79 MXC-ME MXC-MO MXC-PU MXC- 1–1.4 mm wide; margins entire; apex acute or short
TL MXE-GU MXE-HI MXE-QU MXG-VC MXS-MI acuminate; leaf colour deep green or blue-green
on the smooth abaxial face, white stomatal lines
on both adaxial faces. Pollen cones crowded at the
Conservation
proximal end of a new shoot, spreading, subtended
IUCN: NT by light brown bracts, ellipsoid, 6–10 mm long, yel-
low (rarely red), maturing to brown. Microsporo­
 phylls peltate, the distal part cordate to rounded, a­ ltitudes of between 1600 m and 2400 m a.s.l., in the
smooth, ca. 1 mm wide. Seed cones subterminal, south between 2900 m and 3700 m. Stands of this
solitary or frequently in pairs, nearly sessile, falling pine are very open and occur on dry, rocky, exposed
after seed dispersal. Immature cones erect, ovoid, high slopes and ridges, usually devoid of other sig-
dark purple, maturing in two seasons. Mature cones nificant vegetation. Stands may be pure or mixed
ovoid-cylindric becoming more ovoid when open, with P. albicaulis, sometimes Juniperus occidenta­
6–10 × 4–6 cm. Apophysis on seed scales thick, tri- lis grows with it, too. Regeneration and growth are
angular to rhombic, transversely keeled, recurved, extremely slow and stands commonly look as if en-
red-brown. Umbo dorsal, transverse-triangular tirely composed of veteran trees of great age. Regen-
at base, unarmed or terminating in a weak prickle eration is probably episodal and may be linked with
638 1 mm long. Seeds ellipsoid or narrowly obovoid, climatic cycles. Unlike its even longer lived ‘cousin’
8–10 × 4–6 mm, light brown with darker red-brown P. longaeva, growing only 35–40 km to the east of the
specks. Seed wings 10–12 mm long. Sierra Nevada, little is known about the exact ages
of some of the oldest trees, but they are likely to be
more than 2000 years old.
Taxonomic notes
The two disjunct populations have been treated as
Conservation
distinct subspecies or varieties, with the southern
population named P. balfouriana subsp. austrina. This species may be at risk in the long term from
The differences are very small and include chemi- climate change if this is continuing to accellerate,
cal characters (terpene compounds), which are of possibly bringing in competitors and/or pathogens
doubtful significance in taxonomy. The populations it may not be able to cope with. At present, both
being disjunct, it is likely that some differences can disjunct populations are well protected within Na-
be found, e.g. in leaf colour, and that these may be tional Parks and National Forest Wilderness Areas
partly genetic in origin. Mostly the differences in- and ­unaffected by long-term effects of fire suppres-
volve character gradients and would appear to be sion in forests as the trees usually occur in remote
not truly distinct if wide sampling was done be- subalpine locations where such measures have not
fore analysing the data. Pinus balfouriana appears been undertaken. It would be profitable to study the
to be closely related to P. longaeva, a species which (aut-)ecology of this species in more detail in order
occurs to the east, also in isolated populations on to be able to estimate risks under various climate
high mountains. There are at least some consistent change scenarios.
differences in the morphology between these two IUCN: NT
species, but it could as well be defended that there
is only a single species with two or three subspecies
Uses
or ­varieties.
Foxtail pine is not a timber tree due to extremely
slow growth and general inaccessibility of the stands
Distribution
of relatively small trees. The dense and hard wood
USA: mainly California, in two disjunct populations is obviously of value for special uses like wood craft
in northern (Klamath Mts.) and in southern (Sierra and some dead and down wood may be used for this
Nevada) California. One locality (metapopulation) purpose. Almost all stands of this pine are now with-
of the northern population is just across the state in protected areas and felling as well as dead wood
border in Oregon (Lanner, 2007). collecting are there strictly prohibited. This species
TDWG codes: 73 ore 76 CAL was introduced to Britain in the early 1850s by the
Scottish ‘Oregon Association’ through the serv-
ices of John Jeffrey, who collected seeds of western
Ecology
American trees for the gentlemen who instituted the
Pinus balfouriana occurs in the subalpine to alpine association for this purpose. It is still in cultivation
zones of the Klamath Mountains (the northern for gardens, but rare and virtually restricted to col-
population) and of the southern Sierra Nevada (the lections of trees and shrubs. Only two cultivars have
southern populations). In the north it is found at been named, both in the USA.
Pinus banksiana Lamb., Descr. Pinus 1: 7, t. 3. 1803. TDWG codes: 70 NUN NWT-MK 71 ABT BRC MAN
Type: Illustration in Lambert, Descr. Pinus 1: t. 3. SAS 72 NBR NSC ONT QUE 74 MIN WIS 75 INI MAI
1803. (lectotype). MIC NWH NWY

Etymology Ecology
This species has been named after Joseph Banks Pinus banksiana is a boreal to subarctic pine with a
(1743–1820), the famous English naturalist-explorer wide distribution in the lowlands of the North (max.
and President of the Royal Society, who collected alt. 800 m a.s.l.), where it reaches the Arctic tree line
this species in Newfoundland in 1766. and merges with the tundra. Except for a small part
of its range in Nova Scotia, it occurs in continental 639
Vernacular names climate conditions with short, warm summers, and
long, very cold winters and low precipitation, about
Jack pine, Black pine, Hudson Bay pine
half of it as snow. It grows mostly on dry, sandy soils
but is also found on thin soils over granite or met-
Description
amorphosed rock and on peat. This pine is highly
Trees to 27 m tall, but usually smaller and sometimes adapted to fires, which are frequent in the taiga for-
a shrub; trunk to 60 cm d.b.h. Bark scaly, with shal- ests and can destroy vast areas of forest cover. Its se-
low fissures, orange- to red-brown, weathering dark rotinous cones rarely open without the heat of fire
grey-brown. Branches spreading or descending, and remain for many years on the branches. Jack
with higher order branches often pendulous, form- pine is the most successful coniferous pioneer after
ing a pyramidal or irregular to bushy crown. Shoots fire, producing seeds at an early age. It is often ac-
smooth and glabrous, slender, orange-brown or red- companied by Betula papyrifera or Populus tremu­
dish brown; terminal bud ovoid, red-brown, resin- loides. Later successional phases may bring in Picea
ous, 0.5–1 cm long; lateral buds smaller. Cataphylls mariana especially in boggy situations, and Larix
with nearly entire margins. Leaves in remote fasci- laricina, Picea glauca and Abies balsamifera. On the
cles of 2 held by a 3–6 mm long, persistent sheath southern line of its distribution Pinus banksiana can
of perular scale remnants, spreading or ascending, be a component of a more diverse mixed forest with
persisting 2–3 years, often curved and twisted and Pinus resinosa and broadleaved trees such as Quer­
stiff, 2–4.5(–5) cm long, 1.5–2(–2.5) mm wide, light cus spp. and Acer rubrum.
green or sometimes dark green; margins minutely
serrulate; apex acute; stomata in fine lines on all
Conservation
surfaces. Pollen cones in small clusters, spirally ar-
ranged, cylindric, 10–15 mm long, yellow turning IUCN: LC
orange-brown. Seed cones in pairs or whorls of 2–4
(sometimes solitary) at intervals on branches, long
Uses
persistent, opening soon or more often serotinous,
sessile or on short stalks, oblong, 3–5.5 cm long, Despite its relatively small size, Jack pine is an im-
asymmetric, when closed strongly curved inwards portant timber tree for pulpwood, lumber and round
towards the branch, with a tapering apex. Seed scales timber, mainly because it is the most widely distrib-
with generally flat or depressed, but in some cones on uted species of pine in Canada and its pioneer ecol-
the exposed side nearest base more or less gibbous, ogy guarantees even-aged stands with high yields
light brown to tan apophyses; umbo central, small, per ha even under natural conditions. Its wood is
depressed or sunken, unarmed or with a minute, also used in carpentry and joinery, for containers,
reflexed prickle. Seeds obovoid, flattened, 4–5 mm pallets and crates as well as particle-board. It was in-
long, brown to blackish, with a 10–12 mm long wing. troduced to Europe in 1785, but has limited value in
horticulture and is present only as specimen trees
in most arboreta in countries or regions with a cool
Distribution
to cold climate. A few mutants, some from witches
N North America, from Nova Scotia and Pennsylva- brooms, are grown as cultivars. Extensive planting
nia to Northwest Territories and Alberta. in forestry has usually been associated with affores-
 tation of poor sandy soils in northern regions; there usually slightly curved, 12–20 cm long, 3–4 cm wide
it remained a small tree of irregular shape in most when closed and 5–7 cm wide when mature and
instances. opened, usually resinous, soon falling with peduncle
after seeds have been released. Seed scales cuneate-
oblong, widest just below the apophysis, thin woody,
Pinus bhutanica Grierson et al., Notes Roy. Bot. only slightly flexible, with two seed cavities near
Gard. Edinburgh 38: 299. 1980. Pinus wallichiana adaxial base, dull red-brown with lighter marks
A. B. Jacks. subsp. bhutanica (Grierson et al.) of seed wings. Apophysis rhombic, slightly raised,
Businský, Acta Pruhoniciana 68: 10. 1999. Type: obtusely keeled, light brown with darker terminal
Bhutan: Mongar District, Tashiyangsi, 4 km N of umbo. Seeds obovoid, 6–8 mm long, 4–5 mm wide,
640 Tschilingor, A. J. C. Grierson & D. G. Long 2282 slightly flattened, brown; wing persistent, 15–22 mm
(holotype E). long, 7–10 mm wide, grey-brown.

Etymology Taxonomic notes

The species epithet refers to Bhutan, from where it This species was formerly not recognized as distinct
was first described. from Pinus wallichiana and herbarium specimens
belonging to P. bhutanica may remain unrecognized
as the distinction is more in the habit (strongly pen-
Vernacular names
dulous leaves) and young shoots (not always col-
Bhutan pine, Bhutan White pine lected). Some botanists regard it as a subspecies or
variety of Pinus wallichiana. The identification of a
seedling in China is therefore dubious, as seedling
Description
characters were not originally described from this
Trees to 25 m tall or more; trunk to 80 cm d.b.h., species and no comparative study growing both spe-
bole straight and columnar. Bark smooth on young cies has been published.
trees and on branches of large trees, becoming fis-
sured and breaking into small scales that flake off in
Distribution
small chips, dark greyish brown. Branches whorled,
spreading wide and sinuous, those of higher orders Bhutan; NE India: Arunachal Pradesh (Kameng dis-
drooping but with upturned ends, forming a some- trict); SW China: NW Yunnan, SE Xizang [Tibet].
what open crown. Foliage branches slender or stout, TDWG codes: 36 CHC-YN CHT 40 EHM-AP EHM-
new shoots glandular pubescent, whitish pruinose, BH
becoming pale green and then grey. Buds ovoid-
conical; terminal bud 10–15 mm long; lateral buds
Ecology
smaller and more ovoid, slightly resinous; cataphylls
grey with an orange tinge or light reddish brown. From the data on herbarium labels this species ap-
Leaves in fascicles of 5, held in deciduous, basal fas- pears to occur in a variety of forest types, includ-
cicle sheaths of orange-brown flimsy scales, persist- ing mixed pine forest, mixed oak-pine forest, mixed
ing 1.5–2 years, very slender and flexible, 15–24 cm broad-leaved forest, secondary forests of these types,
long, pendulous, often with a bend near the base including thinned (managed?) forest. The elevation
but otherwise straight, 1 mm wide, triangular in (from GIS) is 729–2613 m, from herbarium speci-
cross-section; margins minutely serrulate; leaf col- mens 1676–2750 m a.s.l. (the latter probably more
our bright green abaxially, glaucous white adaxially; reliable in this case).
stomata in fine lines on the two adaxial faces. Pollen
cones in small clusters at base of new shoots, spirally
Conservation
arranged, with 8–12 basal perular scales remain-
ing and partly covering the short cylindrical, yel- The known range of this species (extent of occur-
low cones. Seed cones solitary or in whorls of 2–6, rence or EOO) does not meet the threshold for a
initially erect on stout, (1–)4–6 cm long peduncles, threatened category. It is also thought to occur more
becoming pendulous when growing, cylindrical, widely eastwards, but botanical collecting has been
virtually absent in this region since this species was held by a 10–16 mm long, persistent sheath, retained
first described in 1980. It is reported to be harvested on branchlet 2–3 years, straight and rigid, spreading,
as part of a mixed timber resource, but there is no rarely lax and pendulous, (5–)10–18(–29) cm long,
evidence of decline and new localities have been 1–1.5 mm wide; margins minutely serrulate; leaf col-
found. New collections have recently been made in our bright or dark green; stomata in fine lines on all
China (Yunnan, Gaoligong Mts.) and an assessment surfaces. Pollen cones spirally arranged, short cylin-
of the species in this area is required. drical, 1–2 cm long, yellow. Seed cones solitary or in
IUCN: LC whorls of 2–3(–4), short pedunculate to nearly ses-
sile, persistent, spreading forward or at right angle to
shoot when full grown, narrowly or broadly ovoid-
Uses
conical or ovoid to rarely subglobose when closed, 641
The uses as a timber tree are unknown. In horticul- (4–)6–11(–13) cm long, variously serotinous, 3–5
ture, Bhutan pine has been introduced in the UK cm wide when closed, 5–8 cm wide when opened.
in 1979 and it now grows well in S England, as well Seed scales thick woody, rigid, straight, oblong; apo­
as in Ireland (Grimshaw & Bayton, 2009: 590–591). physes nearly flat or slightly raised, (sharply) trans-
However, it is still restricted to botanic gardens and versely keeled and with thin rays radiating from the
arboreta despite its appealing habit and gracefully centre, more or less rhombic or often with a rounded
pendulous foliage. upper margin, to 20 mm wide at mid-cone, lustrous
red-brown weathering grey; umbo flat or depressed,
4–7 mm wide, broadly rhombic in outline, tan or
Pinus brutia Ten., Fl. Napol. 1, Prodr.: lxxii. 1811–15. grey-brown, unarmed. Seeds obovoid, slightly flat-
tened, 6–7(–8) mm long, grey-brown, sometimes
dark mottled; wing 14–20 mm long, 8–11 mm wide,
Etymology
oblique, grey-brown with darker streaks.
The species epithet is presumably derived from Bru-
tium, a Roman district, now Calabria in southern
Distribution
Italy.
E Mediterranean Region; around the Black Sea;
Caucasus; Turkey; NW Iran; N Iraq.
Vernacular names
TDWG codes: 13 GRC KRI TUE 14 KRY 33 NCS-KR
Calabrian pine, Brutia pine; kizilcam (Turkey) TCS 34 CYP EAI IRN IRQ LBS-LB LBS-SY TUR

Description Ecology
Trees to 30(–35?) m tall; trunk to 1.5(–2.1) m d.b.h., Pinus brutia can form extensive, relatively open
usually a straight bole, or slightly sinuous, some- pine forests, either pure or mixed with Cupressus
times forked. Bark thin, orange-brown, becoming sempervirens and Juniperus excelsa, or mixed open
thick only on lower trunk of large trees, then deeply woodland with Quercus coccifera or Q. calliprinos,
fissured longitudinally, scaly, breaking into elon- Pistacio lentiscus and other drought tolerant trees. It
gated plates, pale brown to red-brown. Branches regenerates after fire by seed dispersal and can suc-
long, spreading and ascending, forming a broad py- cessfully invade maquis vegetation when this does
ramidal or rounded, open crown. Foliage branches not burn for several years. The near-coastal natural
slender, 3–5 mm thick (to 10 mm on cone-bearing distribution of this pine coincides with the Mediter-
shoots), becoming rough with pulvini from fallen ranean climate characterized by cool, wet winters
leaf fascicles, glabrous, new shoots glaucous green, and hot, dry summers. In contrast with planted for-
yellowish brown and finally grey. Buds ovoid-con- ests, natural forests of Pinus brutia have a diverse
ical, acute, 10–15 mm long, not resinous; cataphylls undergrowth of shrubs and herbs and form impor-
with recurved apices, reddish brown fringed with tant habitat for wildlife. The altitudinal range of this
white hairs. Leaves in fascicles of 2 (sometimes 3), species is from near sea level to 1500 m.
 Pinus brutia Ten. var. eldarica (Medw.) Silba,
Uses
Phytologia 58: 367. 1985. Pinus eldarica Medw.,
Pinus brutia has been planted extensively in coun- Trudy Tiflissk. Bot. Sada 6 (2): 21. 1903. Type:
tries around the eastern Mediterranean and Black Illustration in Trudy Tiflissk. Bot. Sada 6 (2): 21.
Sea as it is the easiest pine to grow (with P. halepensis) 1903 (lectotype).
in the Mediterranean climate. It was originally de-
scribed from Calabria in Italy, which is probably a
Description
planted source. Frequent use of P. halepensis sources
from the western Mediterranean threaten to destroy Seed cones ovoid-globose; apophyses slightly raised,
the genetic distinctions between the two species, whitish grey.
642 possibly also in natural stands of P. brutia. The latter
species has a ‘better’ stem shape and growth from a
Taxonomic notes
forestry point of view and should therefore be pro-
tected. Its timber is used for fencing posts, telephone Caucasian and Russian botanists usually refer to
posts, building timbers, railway sleepers, carpentry, this taxon as a distinct species, Pinus eldarica; how-
boxes and crates, hardboard and pulp. The resin of ever, its distinctive characters are minor and may
both pines has been used from ancient times to fla- not warrant taxonomic recognition at all. A critical
vour white wines known as retsina and is still tapped taxonomic revision involving the species pair P. bru­
especially in Turkey, now mainly for the production tia – P. halepensis is overdue and should include this
of turpentine. In horticulture its use is less com- taxon. The use of DNA sequence analysis in such a
mon, mainly as an occasional amenity tree in vil- study is paramount, as morphology is not very in-
lages and towns around the Mediterranean Sea; this formative in these species.
species was also tried as a forestry plantation tree in
SE ­Australia.
Distribution
4 varieties are recognized: Afghanistan?; Azerbaijan (near border with Geor-
gia); Georgia; NW Iran; N Iraq.
Pinus brutia Ten. var. brutia. Pinus halepensis TDWG codes: 33 TCS 34 IRN IRQ
Mill. var. brutia (Ten.) A. Henry, Trees Great
Brit. Ireland 5: 1100. 1910; Pinus halepensis Mill.
Ecology
subsp. brutia (Ten.) Holmboe, [Stud. Veg. Cyprus]
Bergens Mus. Skr., ser. 2, 1 (2): 29. 1914. Type not Semi-dry pine forests, usually mixed with sclero-
designated. phyllous angiosperms in an open canopy.

Pinus brutia Ten. var. densifolia F. Yaltirik & M. Boy-

Conservation
dak, Karaca Arbor. Mag. 5 (4): 175. 2000.
The range of this variety of P. brutia remains imper-
fectly known and an assessment of its conservation
Description
status is also made difficult for lack of information
Leaves 10–18 cm long, spreading. Seed cones ovoid- about trends at (sub)population level.
conical; apophyses red-brown. IUCN: DD

Pinus brutia Ten. var. pendulifolia Frankis, in

Distribution
Taskin (ed.), Papers Int. Symp. Pinus brutia Ten.:
E Mediterranean Region; Turkey. 14. 1993. Type: Turkey, Mugla, near Kayadibi,
TDWG codes: 13 BUL GRC KRI TUE 34 CYP EAI M. P. Frankis 63 (holotype E)
LBS-LB LBS-SY TUR
Description
Conservation
Leaves 18–29 cm long, pendulous. Seed cones ovoid-
IUCN: LC conical; apophyses red-brown.
 multi-coloured pattern on trunk and large branches.
Distribution
Branching ascending or assurgent, forming a broad-
Turkey, Mugla Prov. ly domed or flat-topped crown in older trees. Foli-
TDWG codes: 34 TUR age branches slender, smooth, glabrous, grey-green.
Buds ovoid, to 10 mm long, not resinous; cataphylls
imbricate, red-brown. Leaves in remote fascicles
Conservation
of 3, held by a deciduous basal sheath, spreading,
IUCN: LC straight, rigid, 6–10 cm long, smooth, up to 2 mm
wide, broadly triangular in cross-section; apex acute,
Pinus brutia Ten. var. pityusa (Steven) Silba, leaf colour light green or slightly glaucous; stomata
Phytologia 58: 367. 1985. Pinus pityusa Steven, Bull. on all surfaces. Pollen cones in elongated clusters on 643
Soc. Imp. Naturalistes Moscou 11: 49. 1838. Type not new shoots, spirally arranged, axillary to broad cata-
designated. phylls, ovoid-cylindrical, 10–15 mm long. Seed cones
solitary on short peduncles or nearly sessile, erect
Pinus pityusa Steven var. stankewiczii Sukaczev, Bot. or spreading, broadly ovoid to subglobose when
Zurn. (St. Petersburg) 1: 37. 1906; Pinus brutia Ten. open, 5–7 cm long, 4–5(–6) cm wide, turning from
var. stankewiczii (Sukaczev) Frankis, in Taskin (ed.), pale green to light brown. Seed scales relatively few,
Papers Int. Symp. Pinus brutia Ten.: 14. 1993. broadly cuneate-oblong, woody and rigid. Apophy-
sis much thickened, more or less rhombic in outline,
strongly transversely keeled; umbo dorsal, more or
Description
less central, broadly triangular, armed with a rigid,
Leaves 5–8 cm long, rigid and spreading. Seed cones curved and sharp prickle or spine. Seeds obovoid,
ovoid-conical; apophyses red-brown. 8–10 × 5–6 mm, dull brown, with a reduced wing of
only 4–5 mm which easily detaches from the seed.
Distribution
Caucasus: Krasnodar, Transcaucasia (Abkhazia); Distribution
Georgia; Ukraine: Krym [Crimea].
China: S Gansu, S Hebei, W Henan, Hubei (Badong
TDWG codes: 14 KRY 33 NCS-KR TCS-AB TCS-GR
Xian), Shaanxi, W Shandong, Shanxi, N Sichuan.
TDWG codes: 36 CHC-HU CHC-SC CHN-GS CHN-
Conservation
HB CHN-SA CHN-SD CHN-SX CHS-HE
IUCN: VU (A1c+2c)
The geographic range given above is based on the
mapping of herbarium specimens, mostly in Beijing
Pinus bungeana Zucc. ex Endl., Syn. Conif.: 166.
(PE)*, which we believe have been collected from
1847. Type not designated. Fig. 211, 212
trees growing in the wild. This map is roughly in
agreement with that given by Ying et al. (2003). Oth-
Etymology
er sources mention a wider distribution but seem to
This species was named after the Russian botanist include planted specimens as well. This species is
Alexander Andrejewitsch von Bunge (1803–1890). highly ornamental and has been planted in temple
grounds etc. for many centuries.
Vernacular names
* courtesy Dr. Qiaoping Xiang, Institute of Botany,
Lace-bark pine; bai pi song (Chinese)
Chinese Academy of Sciences, Beijing.

Description
Ecology
Trees to 25–30 m tall; trunk monopodial or multi-
stemmed from a short length above base, to 3 m Pinus bungeana occurs in mountains, often on lime-
d.b.h. Bark smooth, hard, irregularly exfoliating stone rocks and on S-facing slopes, scattered and
with thin flakes, exposing light patches, which later mixed with P. tabuliformis and angiosperms. In the
turn from yellow-green to purplish brown, creating a northern part of its range it is also common on acid
 soils. Its altitudinal range is probably between 500 breaking into large, irregular plates divided by deep
m and 2150 m a.s.l.; records from lower elevations fissures, exfoliating in smaller chips exposing pat-
are probably not from natural occurrences, but from terns of grey, white, purplish and red-brown colours.
planted trees. It is a light demanding species and Branches spreading and assurgent, lower branches
therefore is usually restricted to sites less suitable for curving down, sometimes slowly or poorly self-
other trees, especially angiosperms. pruning, forming rounded or in old trees domed
to flat-topped crowns. Foliage branches slender or
stout, glabrous, rough with pulvini from fallen leaf
Conservation
fascicles, yellowish green becoming brown-grey.
IUCN: LC Buds large, the terminal bud to 3.5 cm long, ovoid-
644 conical, acute, scarcely resinous; cataphylls lanceo-
late, orange-brown, with white fringed margins and
Uses
reflexed apex. Primary leaves on seedlings short,
Lace bark pine is a decorative tree because of its lax, bluish, long retained and often re-appearing on
patterned bark reminiscent of plane (Platanus), epicormic shoots in mature trees. Leaves in fasci-
but more colourful (although in old trees becom- cles of 3, held in persistent, 15–20 mm long sheaths,
ing nearly white) and it is commonly cultivated in retained for 2 years, forming short but dense tufts
China as an amenity tree. It has traditionally been at ends of upturned branches, spreading or slightly
planted around temples and in cemeteries in north- drooping, 20–30 cm long, pliant, 1.5–2 mm wide,
ern China and Korea, from where some of the ear- bright green; margins serrulate; apex acute; stomata
lier records of the species originate, including that in fine lines on all faces, more conspicuous on the
by Alexander von Bunge in Beijing in 1831. It was two adaxial sides. Pollen cones numerous in long
introduced to England in 1843 but is still not very clusters along lower part of new shoots, spirally
common in gardens and parks of Europe. Of lesser arranged, subtended by fringed cataphylls, ovoid-­
importance is the use of its timber, but in NE China cylindrical, yellow. Seed cones solitary or with 2–3 in
it is locally a source for roundwood such as poles a whorl on stout peduncles, erect but soon becoming
as well as construction timber, fences and gates, and reflexed or pendant, 10–20(–25) cm long, narrowly
utilities such as boxes, crates and pallets, while more ovoid when closed, broadly conical with a flat base
refined applications are in furniture and veneers. when opened, when falling a few basal scales may
The seeds are edible and are used in traditional Chi- be left on the branch. Seed scales spreading wide,
nese medicine to alleviate respitatory ailments. oblong, thick woody, rigid, dark brown with lighter
seed wing marks. Apophysis prominently raised,
more or less rhombic, strongly transversely keeled,
Pinus canariensis C. Sm., in Buch, Phys. Beschr. sometimes with additional radial ridges, lustrous
Canar. Ins.: 159. 1825. Type: Macaronesia: Canary red-brown or buff; umbo prominent, obtuse, darker
Islands, C. Smith (1) s.n. (lectotype C). Pl. 27 coloured. Seeds obovoid, 12–15 mm long, brown;
wing adnate (persistent), oblique, 30–35 mm long,
dark grey or grey-brown with dark streaks.
Etymology
The species epithet refers to the Canary Islands,
Distribution
where this pine is native.
Macaronesia, Canary Islands (La Palma, Tenerife,
Gran Canaria, El Hierro).
Vernacular names
TDWG codes: 21 CNY
Canary pine, Canary Islands pine; pino canario
(Spanish)
Ecology
In the Canary Islands this species occupies fairly ex-
Description
tensive forest areas above an altitude of 1200 m to
Trees to 45 m tall; trunk to 2.5 m d.b.h., straight ca. 2200 m a.s.l. It grows well on exposed mountain
or slightly curved. Bark becoming thick on trunk, slopes of volcanic origin amidst or on old lava flows
 645

3
4

Pl ate 2 7. Pinus canariensis. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cone.
 and on scoria. Winter snow and frost limit its alti- introduced to Italy and South Africa as a forestry
tudinal range upwards (Pico del Teide is 3718 m); plantation tree. Its wood is suitable for building
at lower elevations a thermophile woodland and/or timbers because of its good strength, and is widely
xerophytic scrub vegetation, interspersed with rare used in light construction, carpentry, wall panelling,
evergreen lauraceous woodland on N-facing slopes turnery, and marquetry. Mass uses as for pit props
and in moist gullies, dominates. Pinus canariensis is in mining and chipboard are now discontinued, as
dominant within this pine belt where it exists, but forests are required to regrow to former extent and
spring-flowering leguminose and other herbs and capacity after a period of overexploitation. In horti-
shrubs may be common. Some common shrubs are culture it is uncommon, but present in gardens and
Adenocarpus foliolosus, Chamaecytisus proliferus, parks in the Mediterranean region and in some ar-
646 and Cistus symphytifolius. The pine woods are char- boreta elsewhere in countries with a similar climate.
acteristically very open and sunny and often there is Growing this and other pines from warm climates
nothing else on the ground than rocks, pine needles under glass in cooler countries will not succeed;
and great numbers of the large seed cones rolled into pines need the movement of wind to build stems
hollows. that can support a crown. Specimens I have seen
in glasshouses beyond sapling size had to be hung
from the ceiling by a steel wire.
Conservation
Extensive logging in the past combined with de-
structive forest fires at one time reduced the area Pinus caribaea Morelet, Rev. Hort. Côte d’Or 1: 107.
of occupancy (AOO) very substantially. However, 1851.
extensive replanting since the 1950s and regrowth
and better management, as well as cessation of ex-
Etymology
ploitation, have restored most of the pine forests,
even if large trees are now very rare. The fire hazard The species epithet refers to its occurrence in the
still remains and may become more acute with the Caribbean; the species was originally described
ever increasing tourism and its ramifications, and from Isla de la Juventud [Isla de Pinos] in Cuba.
with climate change if this tends to lengthen peri-
ods of drought on the islands where this species is
Vernacular names
indigenous. Development, especially on Tenerife, is
encroaching into the pine belt (“corona forestal”), Caribbean pine; pino de la costa, pino colorado,
which once encircled the volcano and mountain pino macho (Spanish)
spur to the NE, but is no longer contiguous.
IUCN: NT
Description
Trees to 20–35(–45) m tall, d.b.h. to 50–100 cm;
Uses
trunk monopodial, erect. Bark rough, scaly, break-
Pinus canariensis was heavily exploited for its tim- ing into irregularly square plates, grey-brown.
ber in the 19th and early parts of the 20th century; the Branches spreading or ascending, of higher orders
wood was exported. Reforestation with this species similar, or drooping. Crown broad conical, open or
has been sucessful, but today the timber is consid- irregular. Shoots multi-nodal, very rough, resinous;
ered less important than the water retaining capac- with large, short decurrent and persistent pulvini.
ity of the belt of pine forest especially on Tenerife Cataphylls 15–20 mm long, subulate, strongly re-
and Gran Canaria. The ever increasing tourism and curved, scarious, with hyaline-ciliate margins, dark
development of built areas put a great demand on brown. Vegetative buds ovoid-oblong to cylindrical;
limited resources of drinking quality water, which terminal bud 20–25 mm long, but lateral buds small-
even with ample precipitation on the western is- er, ovoid-acute, (slightly) resinous. Fascicle sheaths
lands would be quickly lost downhill and into the 15–20 mm long, retaining their length or reduced to
sea through the extensive areas of scoria and cavern- ca. 10 mm at maturity. Leaves in fascicles of (2–)3
ous lava flows, if it were not for the water retaining (–4), very rarely 5, persisting 3 years, straight,
capacity of the pine forest belt. This ­species has been (slightly) twisted, rigid, (12–)15–26(–28) cm long,
(1.2–)1.4–1.8 mm wide; margins serrulate; apex Roo, Mexico, which is the northernmost occurrence
acute-pungent; leaf colour light or dark green (oc- of P. caribaea var. hondurensis. Pinus caribaea has
casionally glaucous green), more or less lustrous. been widely planted as a timber tree in tropical and
Stomata conspicuous on all faces of leaves, in 8–11 subtropical lowland areas around the world. The
(–14) lines on the convex abaxial face and 4–6 lines heavily resinous wood is used for building, crates,
on each adaxial face. Pollen cones cylindrical, often pallets, boat decks, poles and posts, plywood, and
curved when mature, 2–3 cm × 5–6 mm, pink or yel- particleboard, as well as for pulp manufacture. The
low, turning yellowish or reddish brown. Seed cones latter use comes primarily from plantations outside
subterminal, mostly in pairs or whorls of 3–5(–8), its natural range. In Honduras, trees have been wide-
on 2–2.5 cm long peduncles, ovoid-conical or ovoid ly tapped for resin, but this activity has become less
when opened, more or less symmetrical but base common recently. This species is scarcely known in 647
obliquely flattened, (4–)5–12(–13) × (3–)4–6(–7) cm horticulture and can only be grown successfully in
when open. Seed scales oblong, straight or recurved, the subtropics and tropics.
dark (purplish) brown or blackish brown. Apo­
physis raised or nearly flat on basal scales, rhombic 3 varieties are recognized:
to pentagonal in outline, transversely keeled, up to
15 mm wide, ochraceous, light or dark brown, lus- Pinus caribaea Morelet var. caribaea. Type: Cuba:
trous. Umbo dorsal, flat or slightly raised or pyrami- Isla de la Juventud [Isla de Pinos], Santa Barbara,
dal and curved upward, with a minute, persistent H. A. Lückhoff 11608 (neotype PRF).
prickle. Seeds obovoid, slightly flattened, 5–7 × 2.5–
3.5 mm, light grey-brown with dark spots, or dark
Description
brown to blackish. Seed wings articulate or adnate,
obliquely ovate or oblong, 10–20 × 5–8 mm, thin, Fascicle sheaths 15–20 mm long, retaining their
semi-transparent, yellowish grey or light brown length at maturity. Leaves in fascicles of 3(–4), very
with dark stripes. rarely 2 or 5, (13–)15–26 cm long, (1.2–)1.4–1.8 mm
wide, light or dark green. Seed cones (4–)5–10(–12)
× (3–)4–6(–7) cm when open; apophysis rhombic to
Distribution
pentagonal in outline, ochraceous or light brown,
Caribbean: Bahamas, Turks-Caicos Islands, W Cuba lustrous. Cotyledons (4–)6–7(–9), 12–25 mm long.
(including Isla de la Juventud [Isla de Pinos]); Mex- Seedlings with an elongated stem; primary leaves
ico (S Quintana Roo); N Guatemala; Belize; Hondu- green, more or less ascending, soon replaced by sec-
ras (including Islas de la Bahía); Nicaragua. ondary leaves.
TDWG codes: 79 MXT-QR 80 BLZ GUA HON NIC 81
BAH CUB TCI
Distribution
Caribbean: W Cuba, Pinar del Río, Isla de la Juven-
Ecology
tud [Isla de Pinos].
See under varieties. TDWG codes: 81 CUB

Uses Ecology
Pinus caribaea, and especially the Mesoamerican Forming pure, open, dry fire-climax forest or open
variety hondurensis, is an important timber tree woodland with undergrowth of grasses or scattered
forming easily accessible, extensive stands in level shrubs on sandy or gravelly, well-drained, acidic
to moderately hilly terrain. As such it is heavily ex- soils. Altitudinal range from 1–700 m a.s.l., with
ploited and although not in danger of extinction in most extensive stands below 400 m. The growing is
any of its major areas, many local and especially out- season continuous in a warm tropical climate with
lying populations which are likely to be genetically long dry spells. Annual precipitation varies mainly
distinct are now severely degraded (Dvorak & Don- with altitude, between ca. 1000–1800 mm, with a
ahue, 1992). One of these is Ejido Caobas, Quintana winter dry season. Frost does not occur.
 cornis) has in recent years led to serious dieback
Conservation
affecting 90% or more of the three subpopulations
IUCN: VU (A1c+2c) on these islands (Martin Hamilton, RBG Kew, pers.
comm. 2006). In 2007, a conservation programme
Pinus caribaea Morelet var. bahamensis (Griseb.) has been started, using apparently resistant indi-
W. H. Barrett & Golfari, Rev. Hort. Côte d’Or 1: 107. vidual trees that have been found growing among
1851. Pinus bahamensis Griseb., Fl. Brit. W. I.: 503. infested and dead trees. The infestation has not yet
1864. Type: Bahamas: New Providence, L. Golfari occurred in the Bahamas and was probably caused
77571 (neotype BAB). by importing infested young cut pines from the USA
as Christmas trees in the late 1990s.
648 IUCN: VU [A2b,c,e B2b(v),c(iv)]
Description
Fascicle sheaths 15–20 mm long, reduced to ca. 10 Pinus caribaea Morelet var. hondurensis (Sénécl.)
cm at maturity. Leaves in fascicles of 3(–2), on Grand W. H. Barrett & Golfari, Caribbean Forest. 23: 65.
Bahama 2–3, (13–)15–26 cm long, (1.2–)1.4–1.8 mm 1962. Pinus hondurensis Sénécl., Conif.: 126. 1868.
wide, light or dark green. Seed cones (4–)5–10(–12) Type: Belize: El Cayo, San Pastor Pine Ridge,
× (3–)4–6(–7) cm when open; apophysis rhombic to Chiquibul F. R., Angel’s Drive, W. H. Barrett 77582
pentagonal in outline, ochraceous or light brown, (neotype BAB).
lustrous. Cotyledons (4–)6–7(–9), 12–25 mm long.
Seedlings with an elongated stem; primary leaves
Description
green, more or less ascending, soon replaced by sec-
ondary leaves. Fascicle sheaths 15–20 mm long, retaining their
length at maturity. Leaves in fascicles of 3 (rarely 2,
4, very rarely 5), (12–)16–28 cm long, (1.2–)1.4–1.8
Distribution
mm wide, light green, occasionally glaucous green.
Bahamas: on Grand Bahama, Great Abaco, New Seed cones (4–)5–12(–13) × 3.5–7 cm when open;
Providence and North and South Andros Islands in apophysis rhombic to pentagonal in outline with
the north; on Great Inagua Island in the south. Also upper margin irregularly undulate, chestnut-brown,
on the Turks-Caicos Islands in three distinct sub- lustrous. Cotyledons 5–8, 20–35 mm long. Seedlings
populations at Pine Cay, North Caicos and ­Middle with an elongated stem; primary leaves glaucous,
Caicos. more or less spreading, development of secondary
TDWG codes: 81 BAH TCI leaves delayed.

Ecology Distribution
On flat, eroded limestone rock with pockets of sandy Mexico (S Quintana Roo); Belize; N Guatemala;
soil (pH 7), on sandy spits and old beaches, forming Honduras (including the Islas de la Bahía); Nicara-
pure, open fire-climax stands, usually with scattered gua.
or dense undergrowth of shrubs (often Sabal pal­ TDWG codes: 79 MXT-QR 80 BLZ GUA HON NIC
metto); also invading open scrubland forming sec-
ondary forest. Altitudinal range 1–10 m a.s.l. Annual
Ecology
precipitation decreases from ca. 1500 mm on Grand
Bahama to ca. 1000 mm on Andros in the northern Mainly distributed on the lowland coastal plains
Bahamas, mean annual temperature is ca. 25 °C and within the Atlantic climatic influence, from the edges
frost does not occur. Hurricanes are a recurring de- of mangrove swamps to lower upland “bunch-grass/
structive factor. pine savannas”. It occurs on well-drained, sandy or
gravelly, acidic soils, forming pure (or sometimes
mixed with P. oocarpa and/or P. tecunumanii) fire-
Conservation
climax forest, or pine-oak forest, with undergrowth
In the Turks-Caicos Islands an epidemic of an acci- of grasses, Pteridium aquilinum, or Sabal palmetto
dentally introduced scale insect (Toumeyella parvi­ nearer the coast, possibly replacing broad-leaved
rainforest under human influence in many areas. may occur on abaxial side. Pollen cones clustered,
The altitudinal range is 1–700(–1000?) m a.s.l. An- spirally arranged, short cylindrical, initially reddish
nual precipitation varies greatly, from the highest turning red-brown. Seed cones intitially erect, be-
measurement at Laguna del Pinar in Nicaragua (ca. coming patent, single or in 2–3 in whorls on very
4000 mm) to the lowest at Los Limones in the inte- short peduncles, remaining closed or opening only
rior of Honduras (ca. 660 mm). Mean annual tem- slightly at maturity, from green or glaucous green
perature is from 21–27 °C and no frosts occur. to purplish when growing, resinous, broadly ovoid,
5–8 cm long, 4–6 cm wide, dark brown when ripe.
Seed scales imbricate, widely cuneate proximal-
Conservation
ly, with 2 deep seed cavities adaxially, soft woody.
IUCN: LC Apophysis broadly rhombic or widely triangular to 649
semi-­orbicular, thickened; margins free or slightly
reflexed; umbo terminal, triangular, obtuse, pale
Pinus cembra L., Sp. Pl. 2: 1000. 1753. Type: Illustra- brown. Seeds oblong-obovoid, 10–16 × 6–8 mm,
tion: “Larix sempervirens, foliis quinis, nucleis eduli­ without a wing or wing rudimentary.
bus” in Breyn in Acad. Caes.-Leop. Carol. Nat. Cur.
Ephem. 7: 8, t. 1. (1719). (lectotype). Fig. 213, 214
Distribution
Europe: Alps, Carpathians.
Etymology
TDWG codes: 11 AUT-AU CZE-CZ CZE-SK GER POL
The species epithet is from the Italian vernacular SWI 12 FRA-FR 13 ITA-IT ROM YUG-BH 14 UKR-MO
name cembro for this pine. UKR-UK

Vernacular names Ecology

Arolla pine, Swiss stone pine; Arve, Zirbel (Ger- In the inner valleys of the Alps Pinus cembra, to-
man); auvier, pin arôle (French); cembro (Italian); gether with Larix decidua, forms open conifer forest
limba (Polish); borovice limby (Slovak) and woodland up to the tree line at between 2200
and 2600 m altitude. Arolla pine may descend down
to 1200 m, where it is usually a rare component of
Description
conifer forest dominated by Picea abies. Centuries
Trees to 25 m tall; trunk to 1.5 m d.b.h., often crook- of intensive grazing mainly with cattle have brought
ed and stunted at high altitude. Bark smooth and the tree line in the Alps down and turned much of
orange-brown on young trees and on branches, the ancient forest into pasture woodland. This veg-
becoming scaly and deeply fissured on trunks of etation is in places dominated by Ericaceae such
large trees, turning grey with red-brown fissures. as Vaccinium myrtillus and Rhododendron ferrugi­
Branches spreading and assurging, forming a broad- neum, grasses and herbs. However, more recently
ly conical crown, often flat-topped in exposed trees. alpine grazing has been substantially reduced and
Foliage branches slender to stout, initially with a the forest is making a come-back on many slopes.
dense, brown pubescence, then glabrous, smooth, Arolla pine is dependent on the European Nutcrack-
light brown. Buds small, ovoid-conical, without res- er (Nucifraga caryocatactes, Corvidae) for its effec-
in; cataphylls reddish brown. Leaves in dense tufts tive seed dispersal and the birds carry the seeds and
towards end of shoots, persiting 2–4 years, in fas- therewith the pines up slope. Pinus cembra is most
cicles of 5 held by deciduous basal sheaths of thin, probably a Siberian element in the European flora
orange-brown scales that fall away in the second and is resistant to –40° frost; unlike accompany-
year, spreading wide or forward, straight or slight- ing the larches, P. cembra has evergreen foliage and
ly curved, more or less rigid to flexible but not lax, it reduces the water content in the needles during
(6–)7–9 cm long, triangular in cross-section, not winter to a minimum. Pinus cembra is slow growing
twisted, 1–1.5 mm wide; margins minutely serru- and can live to great age (>1000 years in the Swiss
late; leaf colour green; stomata in white lines pre- Aletsch­wald) having rot-resistant wood.
dominantly on the two adaxial faces, a few stomata
 Cataphylls small, 2–4 mm long, subulate or trian-
Conservation
gular, acute-acuminate, with erose margins, light
IUCN: LC brown, caducous. Vegetative buds ovoid-oblong to
oval-cylindrical; terminal bud 5–8(–10) ×x 3–5 mm;
lateral buds smaller, not or slightly resinous, light
Uses
brown or ochraceous. Fascicle sheaths short, 4–6
Arolla pine is not a significant timber tree due to mm long, loosely imbricate, the scales (pale) brown,
its slow growth and commonly curved or contort- soon recoiling, then pale straw-coloured to grey,
ed shape, although the tree can grow quite straight forming a small rosette at base of fascicle but decid-
and to considerable size in protected localities. As uous before the leaves fall. Leaves in fascicles of (2–)
650 it grows with European larch and the latter is much 3(–4, rarely 5), persisting (3–)4–5(–7) years, curved
more valued for timber, forestry practises tend to or less often straight, lax or sometimes rigid, (2–)3–
favour the latter, which in succesional terms is the 6(–8) cm long, (0.6–)0.7–1(–1.2) mm wide; margins
pioneer species. The wood has been used for the entire; apex acute-acuminate or pungent; leaf colour
building of traditional houses and is valued for spe- variable, dull green to glaucous green, with or with-
cial uses such as joinery, panelling, cabinet mak- out white adaxial faces; leaves sometimes producing
ing, tools, and wood turning. The seeds, though a few resin drops. Stomata: leaves amphistomatic,
edible, are difficult to harvest due to the soft, res- or epistomatic in one variety, in 2–3(–4) lines on
inous and closed cone scales and are consequently the convex abaxial face (or none), in 2–3(–4) lines
mostly left to the birds. As an ornamental tree it is on each adaxial face. Pollen cones small, ca. 5 × 3
quite valuable, but it is sensible to ‘late frosts’ as are mm, yellowish. Seed cones solitary, paired or more
other conifers from very cold regions where spring rarely in whorls of 3, on very short, 3–5(–8) mm
means spring and is not interrupted by a brief re- long peduncles remaining with fallen cones. Ma-
turn to winter. Despite this, a number of cultivars ture cones seemingly sessile, irregularly globose or
are known, both with distinct habit and with diver- ovoid-globose when closed, spreading often wider
gent needle colours; they are particularly well grown than long when opened, with a flattened base, irreg-
in northern and eastern Europe. ular in size and shape, often resinous, (2–)3–5(–7.5)
× 3–6(–7) cm when open. Seed scales parting eas-
ily and widely, more or less weakly attached to the
Pinus cembroides Zucc., Abh. Math.-Phys. Cl. rachis and hence moveable, up to 15–20 mm wide,
Königl. Bayer. Akad. Wiss. 1: 392. 1832. concavo-convex, with 1–2 deep seed cavities. Apo-
physis raised, transversely keeled or radially keeled,
rhombic to pentagonal in outline but irregular, in all
Etymology
shades from yellowish green or ochraceous to red-
The species epithet means ‘similar but not equal to dish brown, sometimes lustrous. Umbo dorsal, flat
cembra’, i.e. Pinus cembra L. or raised and curved, with a minute prickle. Seeds
obovoid-oblique, 10–16 × 6–10 mm, greyish brown
to blackish grey, or light brown; integument thick,
Vernacular names
0.5–1(–1.1) mm; megagametophyte (‘endosperm’)
Pinyon pine, Mexican nut pine; piñón (Spanish) pinkish or white when fresh. Seed wings absent.

Description Distribution
Shrubs or trees 3–15(–25) m tall, d.b.h. to 10–80 SW USA: SE Arizona, SW New Mexico, SW Texas;
(–120) cm. Trunk monopodial, very short (shrubs) Mexico: from the Sierra Madres to South-Central
to medium size, branching low. Bark thick, rough Mexico and in Baja California (Sierra de la Laguna).
and scaly, breaking into small, irregular, shaggy TDWG codes: 76 ARI 77 NWM TEX 79 MXC-DF
plates, grey or greyish brown. Branches spreading or MXC-ME MXC-PU MXC-TL MXE-AG MXE-CO MXE-
trailing on the ground, or ascending. Crown an open CU MXE-DU MXE-HI MXE-NL MXE-QU MXE-SL
or dense shrub or broad, open, irregular. Shoots or- MXE-TA MXE-ZA MXG-VC MXN-BS MXN-SO
ange-brown or slightly glaucous at first, soon grey.
 Pinus cembroides Zucc. subsp. cembroides var. cem­
Ecology
broides. Type: Mexico: México, Sultepec, Santa Cruz,
In much of its range, Pinus cembroides occupies a F. Karwinski s.n. (holotype M). Fig. 215, 216
transition zone between (semi-)desert plateaux
and valleys and more mesic coniferous montane
Description
forests. The altitudinal range is extensive: (800–)
1500–2600(–2800) m a.s.l., with highest occur- Small tree to 10–15 m tall. Leaves in fascicles of 2–3,
rences in the SE of its range. It grows on a variety of lax or sometimes rigid, (2–)3–5(–6.5) cm × (0.6–)
substrates, ranging from alluvial bajadas to volcanic 0.7–1 mm; leaf colour variable, dull green to glaucous
rock, usually on scarcely developed soils. It forms green especially on adaxial faces. Seeds 10–13 × 6–10
open woodland alone or mixed with Juniperus spp., mm; integument thick, 0.6–1 mm; megagameto­ 651
Pinus nelsonii, P. pinceana, Quercus, Yucca, Agave, phyte pinkish when fresh.
Cactaceae (e.g. Opuntia), Arctostaphylos, Cean­
othus, Arbutus and other shrubs of dry, hot areas; at
Distribution
higher and/or moister sites it forms part of a mixed
pine-oak woodland or forest including, e.g. Pinus SW USA: SE Arizona, SW New Mexico, SW Texas;
arizonica, P. engelmannii, P. leiophylla var. chihua­ Mexico: NE Sonora, Chihuahua, Coahuila, Duran-
huana, and P. pseudostrobus in the southeastern part go, Zacatecas, Nuevo León, W Tamaulipas, San Luís
of its range. The climate is warm and dry, with an- Potosí, Aguascalientes, NE Jalisco, N Guanajuato,
nual precipitation varying from 380–650 mm and a Querétaro, Hidalgo, México, Distrito Federal, Tlax-
dry season of 7–8 months. Frost may occur at higher cala, Veracruz and Puebla.
elevations in the interior, but is infrequent. There is TDWG codes: 76 ARI 77 NWM TEX 79 MXC-DF
an important mutualist relationship with the corvid MXC-ME MXC-PU MXC-TL MXE-AG MXE-CO MXE-
birds Aphelocoma coerulescens and Gymnorhinus cy­ CU MXE-DU MXE-GU MXE-HI MXE-NL MXE-QU
anocephalus (Tomback & Linhart, 1990), which feed MXE-SL MXE-TA MXE-ZA MXN-SO
on the seeds and cache them, thereby providing an
effective dispersal mechanism.
Conservation
IUCN: LC
Uses
Although not a timber tree in most areas due to Pinus cembroides Zucc. subsp. cembroides Little
its low stature and low and heavy branching, Pinus var. bicolor Little, Phytologia 17: 331. 1968. Pinus
cembroides is nevertheless an economically impor- discolor D. K. Bailey & Hawksw., Phytologia 44: 130.
tant species of pine in Mexico. Its principal value for 1979; Pinus culminicola Andresen & Beaman var.
local economies lies in the edible seeds (piñones), discolor (D. K. Bailey & Hawksw.) Silba, Phytologia
which are regularly harvested and marketed. Fur- 56: 490. 1985. Type: USA: Arizona, Santa Cruz Co.,
ther use is made of its wood for carpentry, or some- Madera Canyon, Coronado National Forest, Santa
times for timber where there is no other pine species Rita Mountains, Madera Canyon, E. L. Little 23011
available, as in Baja California Sur. Due to its ad- (holotype US).
aptation to semi-arid environments and extensive
range it is also an important shrub or tree for aspects Pinus johannis Rob.-Pass., Adansonia, sér. 2, 18:
of land management, such as watershed protection, 366. 1978; Pinus culminicola Andresen & Beaman
prevention of erosion, and as a shade tree in agro- var. johannis (Rob.-Pass.) Silba, Phytologia 56: 491.
forestry. Pinyon pines are uncommon in cultivation 1985. Type: Mexico: Zacatecas, Concepción del Oro,
and mostly seen in arboreta and botanic gardens Puerto el Dique, M. F. Robert & J. Passini 5936B
and in some urban landscaping schemes mainly in (­holotype P).
SW USA.
Description
3 subspecies and 2 varieties are recognized:
Shrubs or small trees to 3–12 m tall. Leaves in fas-
cicles of 3, rarely 2, 4 or 5, more or less rigid, 2.5–
 5(–6) cm × 0.8–1.2 mm, dull green to grey-green on compete with oaks is not known. Access roads on
abaxial face, usually glaucous white on adaxial faces, the mountain are few and primitive.
divided by a green midrib. Seeds 10–12 × 6–9 mm, IUCN: VU (A1c)
integument thick, 0.5–1 mm; megagametophyte
white when fresh. Pinus cembroides Zucc. subsp. orizabensis
D. K. Bailey, Phytologia 54: 89. 1983. Pinus
cembroides Zucc. var. orizabensis (D. K. Bailey)
Distribution
Silba, Phytologia 68: 48. 1990; Pinus orizabensis
SW USA: SE Arizona, SW New Mexico; Mexico: (D. K. Bailey) D. K. Bailey & Hawksw., Novon 2:
NE Sonora, Chihuahua, Coahuila, Nuevo León, W 306. 1992. Type: Mexico: Puebla, Soltepec, along
652 Tamaulipas, Durango, Zacatecas, N & W San Luís Hwy. 140, ca. 10 km SW of San Salvador el Seco,
Potosí. D. K. Bailey 83–01 (holotype MEXU). Fig. 217
TDWG codes: 76 ARI 77 NWM 79 MXE-CO MXE-
CU MXE-DU MXE-NL MXE-SL MXE-TA MXE-ZA
Description
MXN-SO
Small trees to 8–10 m tall. Leaves in fascicles of 3(–4,
rarely 2 or 5), more or less rigid, (2–)3–5(–6) cm ×
Conservation
0.7–1.1 mm, dull green to greyish green. Seeds 10–14
IUCN: LC × 6–10 mm; integument thick, 0.7–1.1 mm; mega­
gametophyte pinkish when fresh.
Pinus cembroides Zucc. subsp. lagunae (Rob.-Pass.)
D. K. Bailey, Phytologia 54: 98. 1983. Pinus cem­
Distribution
broides Zucc. var. lagunae Rob.-Pass., Bull. Mus.
Hist. Nat. (Paris), ser. B, Adansonia 1: 64. 1981; Pi­ Mexico: Puebla, Tlaxcala, Veracruz.
nus lagunae (Rob.-Pass.) Passini, Phytologia 63: 337. TDWG codes: 79 MXC-PU MXC-TL MXG-VC
1987. Type: Mexico: Baja California Sur, Sierra de la
Laguna, La Laguna, M. F. Robert 10021 (holotype P).
Conservation
IUCN: NT
Description
Trees to 20–25 m tall. Leaves in fascicles of (2–)3,
very rarely 4, lax, (2.5)4–7(–8) cm × (0.7–)0.8–0.9 Pinus clausa (Chapm. ex Engelm.) Sarg., [Rep.
mm, dull green to greyish green. Seeds 10–16 × 6–10 Forests N. America] US 10th Census 1880, vol. 9:
mm; integument thick, 0.5–0.8 mm; megagameto- 199. 1884. Pinus inops Aiton var. clausa Chapm. ex
phyte pinkish when fresh. Engelm., Bot. Gaz. 2: 125. 1877. Type not designated
(possibly a collection from near Apalachicola,
Florida, USA by A. W. Chapman).
Distribution
Mexico: Baja California Sur (Sierra de la Laguna). Pinus clausa (Chapm. ex Engelm.) Sarg. var. immugi­
TDWG codes: 79 MXN-BS nata D. B. Ward, Castanea 28: 4. 1963; Pinus clausa
(Chapm. ex Engelm.) Sarg. subsp. immuginata
(D. B. Ward) E. Murray, Kalmia 13: 22. 1983.
Conservation
The extent of occurrence (EOO) of this subspe-
Etymology
cies is estimated to be 20,000 ha; it is endemic in
the Sierra de La Laguna. The trees now enjoy some The species epithet means ‘closed’ and refers to the
form of protection, but have been exploited for local seed cones.
use, which is estimated to have led to a decline. As
a small population it is prone to destruction by fires
Vernacular names
if these were extensive or frequent. To what extent,
if any, the pines are dependent on fire in order to Sand pine, Florida spruce pine
 Description Distribution
Trees to 21 m tall; trunk to 50 cm d.b.h., often USA: Alabama (Baldwin Co.), Florida.
stunted and crooked, branching low or sometimes TDWG codes: 78 ALA FLA
multi-stemmed; crown open and irregular or flat-
tened. Bark scaly, breaking into narrow ridges and
Ecology
furrows, grey-brown, in the crown thin and flak-
ing, red-brown. Shoots slender, purplish to reddish This lowland pine from the sandy interior and coasts
brown, often glaucous, glabrous, rough with persist- of Florida occupies the drier areas away from swamps
ent pulvini after leaf fascicles have fallen. Buds ovoid and eutrophic rivers. It is a fire-succesional species,
to cylindrical, 0.5–1 cm long, with or without resin, in particular on infertile white sand dunes and flats 653
red-brown; cataphylls with white, fringed margins. of marine origin and low elevations between 5 and
Leaves in fascicles of 2, held in 4–8 mm long sheaths, 60 m a.s.l. or up to 90 m on the west coast. There
persisting 3–4 years, spreading, (2–)5–8(–10) cm are abundant summer rains while winters are gener-
long, nearly straight or contorted, 1–1.5 mm wide, ally dry, but much of the precipitation drains away
rigid, dark green or light green; margins minutely quickly into the sandy soil. There are two ecotypes
serrulate; apex acute; stomata in inconspicuous lines or ‘races’ of this species, one with mostly seroti-
on all surfaces. Pollen cones in small clusters, spiral- nous cones and another of which the cones open
ly arranged, short cylindric, 10–15 mm long, yellow more readily; these were recognized previously as
maturing to red-brown. Seed cones solitary or in botanical varieties. Pinus clausa is characteristic of
pairs, persistent, nearly sessile or short pedunculate, ‘sand pine scrub’ a distinctive plant community of
serotinous or opening soon in warm sunshine, sym- north-central Florida. The tree layer is dominated
metrical, 4–8 cm long, narrowly ovoid when closed, by P. clausa, with an understorey composed of ever-
broadly ovoid when open. Seed scales thin woody, green shrubs to 3 m tall and in almost total absence
rigid, often purple on the border below the apohysis of a herb layer. Several species of oak (Quercus) and
on adaxial surface, turning brown when old. Apo- small palms (Sabal etonia, Serenoa repens) are most
hysis slightly raised, keeled, angular in outline, with abundant. The ground is often covered by lichens
a small, low pyramidal umbo terminating in a small (Cladina evansii, Cladonia spp.).
prickle or unarmed, colour dull reddish brown when
cones are ripened. Seeds obovoid, slightly flattened,
Conservation
4–6 mm long, pale brown with darker spots or dark
brown, with a narrow wing 15–20 mm long. IUCN: NT

Taxonomic notes Uses

Trees with serotinous cones in central Florida Sand pine is of some value as a timber tree only in
have been described as P. clausa var. immuginata. plantations, where it can give a reasonably high yield
Although there appears to be some geographical for the pulp wood industry. The ‘Chocktawhatchee’
separation of the two forms, serotiny of cones has form of western Florida with short branches and
been found among trees with opening cones and dark green needles is grown as a Christmas tree.
this character alone seems insufficiently distinct to The species is thought to be of use in ‘biomass plan-
give it taxonomic separation as a variety or subspe- tations’ for fuelwood from trials conducted by the
cies. Cone serotiny is clearly an adaptive trait re- USDA Forest Service in the 1980’s. Such use may be-
lated to fire and trees with these cones are probably come more important in the current political drive
strongly selected for in areas with frequent fires. The to reduce dependence on oil in the USA and to in-
phenomenon occurs in a good number of phylo­ crease the contribution of ‘renewables’ to the fuel for
genetically distant (not closely related) species in the motorcars. This species has no significance in horti-
subgenus Pinus. culture.
 Pinus contorta Douglas ex Loudon, Arbor. Frut.
Distribution
Brit. 4: 2292. 1838.
Western North America, from Yukon to Baja Cali-
fornia and S Colorado.
Etymology
TDWG codes: 70 ASK NWT-MK YUK 71 ABT BRC 73
The species epithet contorta means twisted (contort- 74 SDA 76 79 MXN-BC
ed) and probably refers to the often twisted needles.
Ecology
Vernacular names
Pinus contorta occupies a large part of the North
654 Lodgepole pine, Tamarack pine, Shore pine American West with in particular a vast latitudinal
range. It consequently has a wide ecological am-
plitude and grows form near sea level to 3350 m or
Description
perhaps higher and from the relatively mild but cool
Shrubs or trees to 50 m tall; trunk to 1–2 m d.b.h., and rainy Pacific coast to the cold and continental
very straight to contorted. Bark scaly, breaking into interior of the northern Rocky Mountains. Pre-
small plates or longitudinal ridges and furrows, light cipitation consequently ranges from only 250 mm
brown, reddish brown to grey. Branching variable, at low elevations in the interior to 5000 mm along
dependent on environment and to some extent ge- the northern coast. In the interior, Lodgepole pine
netically determined, in dense stands short, self- forms pioneer stands of great density after forest
pruning, usually spreading and contorted, lower fires and can form monotypic stands of great ex-
branches descending. Foliage branches slender, tent, especially on infertile soils. In other sites it is
shoots green, turning orange-brown, rough with associated with many western conifers, most com-
pulvini after leaf fascicles have fallen. Buds ovoid, monly in the north with Picea glauca and mixed
the terminal ones narrow, 10–15 mm long, the lateral with Betula papyrifera or Populus tremula; at higher
ones much smaller, slightly resinous; cataphylls dark altitudes with Tsuga mertensiana, Picea engelmannii
red-brown, with entire or scarious margins. Leaves and Abies lasiocarpa. Further south, the species di-
in more or less remote fascicles of 2 held by short versity increases and in California it is a component
sheaths, persisting 4–8 years, spreading, (2–)3–7(–8) of the mixed conifer forest as well as subalpine coni-
cm long, rigid, usually curved and twisted, 0.7–2 fer woodland and meadows with numerous conifer
(–3) mm wide; margins minutely serrulate; leaf col- species. Here soils are more nutrient rich and fires
our light yellow-green to dark green; apex obtuse, are less frequent, so Pinus contorta does not attain
acute or acuminate; stomata in lines on all surfaces. dominance. As a component of the mixed conifer
Pollen cones in small clusters, spirally arranged, el- forest it can attain 50 m in height, with 1 m d.b.h.,
lipsoid to cylindric, 5–15 mm long, orange-red turn- and live to a considerable age. In other areas, such as
ing red-brown. Seed cones solitary or in whorls of large tracts of Yellowstone Park in Wyoming, Lodge-
2–5, nearly sessile to short pedunculate, spreading pole pine appears to be self-perpetuating as the only
or reflexed, variably persistent and variably seroti- tree species capable of growing in a more dynamic
nous, sometimes remaining on branches and closed environment characterized by frequent fires.
for many years, ovoid but often with an asymmet-
rical base, (2–)3–5(–6) cm long, broadly ovoid to
Uses
nearly globose when open. Seed scales thin woody,
rigid, brown. Apophysis more or less rhombic in Lodgepole pine is a major timber tree in west-
outline, variously raised, sometimes gibbous on the ern North America. Its geographical varieties dif-
sun-exposed side near base of cone, transversely fer apart from minor botanical characters also in
keeled, with a central, depressed, triangular umbo growth performance and maximum size, which
armed with a variable, thin or blunt prickle up to 6 is at least in part due to environmental factors. As
mm long. Seeds obovoid, slightly flattened, ca. 5 mm other conifers, it attains greatest size in Oregon and
long, red-brown with blackish spots or nearly black; California (var. murrayana) with trees over 50 m tall
wing 10–12 mm long. and 2 m diam. In the interior, the tall, thin stems
of densely grown pines provided the ‘lodgepoles’, i.e.
tent poles for the bison-hide covered conical tents asymmetric and recurved on branches, persistent
of the Plains Indians with such famous tribes or ‘na- and variously serotinous; apophyses near base of
tions’ as the Dakota (‘Sioux’) and Blackfeet. Today cone often gibbous; umbo with a small, blunt prickle.
Lodgepole pine is put to all traditional (European)
uses common to pine wood, but mass production Distribution
is for the pulp industry or increasingly the manu-
NW North America, from Yukon to Colorado.
facture of so-called structural particleboard, where
TDWG codes: 70 ASK NWT-MK YUK 71 ABT BRC
chips are glued into boards for interior construction.
SAS 73 COL IDA MNT ORE WAS WYO 74 SDA 76 UTA
Pinus contorta is sometimes planted as a shelter tree
on barren sites, but otherwise uncommon in cultiva-
Conservation
tion; only a limited number of cultivars are known. 655
Hybrids between P. contorta and P. banksiana have IUCN: LC
been generated by foresters in the USA in attempts
to produce trees suitable for plantations. Pinus contorta Douglas ex Loudon var. murrayana
(Balf.) Engelm., in Watson, Bot. California 2: 126.
3 varieties are recognized: 1880. Pinus murrayana Balf., in Murray, Bot. Exped.
Oregon 8: 2. 1853; Pinus contorta Douglas ex Lou-
Pinus contorta Douglas ex Loudon var. contorta. don subsp. murrayana (Balf.) Engelm., Trans. St.
Type: USA: Washington, D. Douglas s.n. (lectotype Louis Acad. Sci. 4: 177. 1880. Type: USA: California,
K). Siskiyou Mts, J. Jeffrey 740 (holotype E). Fig. 218

Description
Description
Leaves 2–5 cm long, rarely longer, slender, 0.7–1.2(–
Trees to 50 m tall; trunk to 1–2 m d.b.h. Leaves
1.5) mm wide. Seed cones solitary or in whorls of
5–8 cm long, 1–2 mm wide. Seed cones solitary or
2–5, asymmetrical near base, persistent and various-
in pairs, mostly symmetrical or nearly so, opening
ly serotinous; umbos armed with a slender prickle
soon and falling soon after seed dispersal; apohyses
to 6 mm long.
transversely keeled but not gibbous; umbo with a
small prickle.
Distribution
NW coast of North America, from S Alaska to Distribution
N California.
USA: California, Nevada, Oregon, S Washington;
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
NW Mexico: Baja California Norte.
TDWG codes: 73 ORE WAS 76 CAL NEV 79 MXN-BC
Conservation
IUCN: LC Conservation
IUCN: LC
Pinus contorta Douglas ex Loudon var. latifolia
Engelm., in Watson, Botany (Fortieth Parallel):
331. 1871. Pinus contorta Douglas ex Loudon subsp. Pinus coulteri D. Don, Trans. Linn. Soc. London 17:
latifolia (Engelm.) Critchf., Publ. Maria Moors 440. 1836. Pinus ponderosa Douglas ex C. Lawson
Cabot Found. Bot. Res. 3: 107. 1957; Pinus divaricata subsp. coulteri (D. Don) E. Murray, Kalmia 12: 23.
(Aiton) Dum.-Cours. var. latifolia (Engelm.) 1982. Type: USA: California, Monterey Co., Santa
Boivin, Naturaliste Canad. 93: 272. 1966. Type: Lucia Mts., Cone Peak, T. Coulter s.n. (holotype
USA: Utah, Uintah Mts., S. Watson [“Clarence TCD). Fig. 219, 220
King’s Exped. 40th Parallel”] 1110 (holotype A).
Etymology
Description
This species was named after Thomas Coulter (1793–
Leaves (4–)5–8 cm long, 1–2(–3) mm wide. Seed 1843), an Irishman who collected the type specimen,
cones solitary or in pairs, 4–6 cm long, strongly a large cone now kept in Trinity College, Dublin.
 Vernacular names Distribution
Coulter pine, Bigcone pine USA: California (Coast Ranges); Mexico: Baja Cali-
fornia Norte.
TDWG codes: 76 CAL 79 MXN-BC
Description
Trees to 15–25 m tall, d.b.h. to 1 m. Trunk mono­
Ecology
podial, straight or curved at base. Bark on trunk
rough, scaly, divided into irregular, longitudinal In California, Coulter pine is prominent in the
plates, dark brown with black fissures. Branches southern California Mixed Conifer Forest, especially
656 spreading horizontally to more or less ascending, at the lower limit of this forest type, where it merges
ultimate branches upturned, forming a broad, irreg- into fire-prone chaparral. In Baja California it is also
ular pyramidal and open crown. Shoots multinodal, a tree of mixed chaparral, together with Quercus
thick, rigid, rough with prominent, decurrent pul- chrysolepis, or growing on granite boulder forma-
vini, light orange-brown or purplish brown, often tions around Laguna Juárez. Its altitudinal range is
glaucous. Cataphylls 15–20 mm long, subulate, re- from 300 m to 2100 m a.s.l. (in Mexico 1200–2150
curved, with caudate apex and erose-ciliate margins. m). It is most commonly found on dry, rocky slopes
Vegetative buds large; terminal bud 15–30 mm long, and ridges, where competition from other trees is
ovoid-acute; lateral buds ovoid, smaller, resinous, minimized. The climate is of a Mediterranean type
reddish brown. Fascicle sheaths thick, remaining ca. with winter rain and long, hot and dry summers.
20 mm long. Leaves in fascicles of 3, persisting 3–4
years, very rigid, straight or curved, slightly twisted,
Conservation
15–25(–30) cm long, 1.9–2.2 mm wide; margins ser-
rulate; apex acute-pungent to subulate; leaf colour IUCN: LC
light green to grey-green. Stomata on all faces of
leaves, with (10–)12–14 lines on the convex abaxial
Uses
face and (4–)5–7 lines on each adaxial face. Pol-
len cones ovoid to cylindric, 2–2.5 cm long, light Coulter pine has no particular commercial value
purplish brown, maturing to orange-brown. Seed as a timber tree and its seeds, although edible, are
cones solitary or occasionally in pairs, or in whorls not harvested for consumption. It is quite frequent-
of 3–4(–5) on bole of young trees; borne on thick, ly represented in parks and arboreta in southern
short, persistent peduncles, holding cone to branch ­Europe and the milder parts of the British Isles, and
for as long as 25 years and retaining a few basal this pine has also been introduced as an amenity tree
scales when it falls. Mature cones ovoid, massive, in Australia and New Zealand. In California it is also
heavy, usually slightly oblique or curved, moder- planted in parks and large gardens, often in small
ately serotinous, 20–35 × 15–20 cm when open, ex- groups. No cultivars are known of this pine and it
tremely resinous. Seed scales flat, thick woody with is apparently rarely grown and sold by horticultural
thin, curved margins, widest towards the apophysis. nurseries. The impressive cones are often collected
Apophysis very strongly developed, on one side of and displayed as curiosities in private houses as well
cone larger than on the other, thick woody, sharply as schools and other public buildings.
transversely keeled, merging into a long, uncinate
umbo, up to 30 mm wide, light yellowish brown
or light caramel coloured. Umbo dorsal, elongate, Pinus cubensis Griseb., [Pl. Wright.] Mem. Amer.
curved, with keeled sides, 25–35 mm long, 10–15 mm Acad. Arts, ser. 2, 8: 530. 1862. Pinus occidentalis Sw.
wide at base, ending in a sharp uncinate claw. Seeds var. cubensis (Griseb.) Silba, Phytologia Mem. 7: 55.
obliquely obovoid, slightly flattened, 10–18 × 7–10 1984. Type: Cuba: E Cuba, C. Wright 598 (lectotype
mm, smooth, lustrous dark brown, turning black- GOET).
ish. Seed wings dolabriform to semi-ovate, 18–30 ×
12–16 mm, orange-brown to dark reddish brown. Pinus maestrensis Bisse, Ciencias (Havana), ser. 10,
2: 2. 1975; Pinus occidentalis Sw. var. maestrensis
(Bisse) Silba, Phytologia 68: 57. 1990.
 Etymology Distribution
The species epithet cubensis refers to Cuba, where it E Cuba: From the Sierra Maestra and the Sierra de
is endemic. Nipe E into the highlands terminating the eastern
part of the island.
TDWG codes: 81 CUB
Vernacular names
Cuban pine
Ecology
This species occurs in foothills and highlands as well
Description
as in ‘pine barrens’ along the coast. Its altitudinal 657
Trees to 25–30 m tall, d.b.h. to ca. 1 m; trunk mono­ range is from 100–900(–1200) m a.s.l. Pinus cubensis
podial, erect. Bark thick, rough, scaly, breaking into forms mostly pure but open stands or is invasive in
irregular, squarish plates divided by deep fissures, or disturbed sites on serpentine or serpentine-derived,
forming vertical ridges between fissures, grey-brown, often ferruginous soils (‘Nipe latosol’ or ‘Nipe clay’),
weathering grey. Branches spreading irregularly, or on alluvial sediment near the coast. In the high-
forming a small, irregular, broadly domed and open lands there is abundant rainfall (1800 mm or more
crown. Shoots more or less multi-nodal, glaucous annually), but a dry winter season is a typical aspect
to pruinose in the first and second years. Cataphylls of the tropical to subtropical climate.
small, subulate, scarious, early recurving, with ciliate
margins, brown. Vegetative buds ovoid-oblong; ter-
Conservation
minal bud 10–15 mm long and lateral buds smaller,
acute, not resinous or sometimes slightly resinous. IUCN: LC
Leaf fascicles initially 9–12 mm long, on adult leaves
reduced to 7–9 mm, persistent. Leaves in fascicles
Uses
of 2(–3), persisting 2–3 years, more or less rigid, or
flexible, straight or slightly curved, (6–)10–15(–18) Pinus cubensis is the only pine occurring in the east-
cm long, 0.8–1.3 mm wide; margins serrulate, acute; ern part of Cuba; no pines occur naturally in Cuba
leaf colour light green, occasionally glaucous green. between eastern Cuba and Pinar del Río in the ex-
Stomata on all faces of leaves, with 5–10 lines on the treme west of the island, and as such the pines are of
convex abaxial face and 4–5(–7) on adaxial face(s). importance economically as a timber source to this
Pollen cones cylindrical, often recurved when ma- part of Cuba. This species is not known to be in cul-
ture, 1.5–2 cm × 5 mm. Seed cones sub-terminal, tivation.
solitary or in pairs, less commonly in whorls of 3–4,
on 1–2 cm long, recurved or straight peduncles, de-
ciduous. Mature cones (narrowly) ovoid to ovoid-at- Pinus culminicola Andresen & Beaman, J. Arnold
tenuate when closed, (broadly) ovoid when opened, Arbor. 42: 437. 1961 Type: Mexico: Nuevo León,
(3.5–)4–7 × (2–)3.5–5 cm when open. Seed scales Galeana, Cerro Potosí, J. H. Beaman 2675 (holotype
parting readily except basal ones, oblong, straight or MSC). Fig. 221
slightly recurved, thin woody. Apophyses more or
less symmetrical around cone, slightly raised, trans-
Etymology
versely keeled, transverse-rhombic to pentagonal
in outline, with curved or irregular upper margin, The species epithet means: ‘growing on the summit’
up to 11 mm wide, radially striate, dark ochraceous (of mountains).
to dull brown. Umbo dorsal, depressed or slightly
raised, the minute prickle early deciduous. Seeds
Vernacular names
obliquely ovoid, flattened, 5–6 × 3–3.5 mm, mottled
grey-brown. Seed wings obliquely ovate-oblong, 12– Potosi pinyon pine; Potosí piñon (Spanish)
16 × 5–6 mm, grey-brown, with a graphite-like tinge.
 Description Ecology
Shrubs 1–5 m tall, diam. of stems to 15–25 cm; mul- The altitudinal range of P. culminicola is 3000–
tistemmed, or very low branched, individuals form- 3700 m a.s.l., which includes the highest ridges of
ing a spreading, in places dense vegetation. Bark these mountains. Its habit is very similar to other
thin, scaly, exfoliating in small, irregular plates, mountain ‘dwarf pines’, e.g. P. mugo in Europe and
grey-brown, soon weathering grey. Branches often P. pumila in NE Asia. Adaptation to blasting, ice- or
prostrate to assurgent, the higher order branches sand-laden winds and a short growing season is re-
assurgent to erect. Shoots short, thick, light brown sponsible for this habit. Soils are mostly rocky and
turning grey. Cataphylls small, narrowly triangu- calcareous. Climatic conditions are not well known
658 lar to subulate; apex caudate; margins erose, light due to a lack of weather stations at these summits, but
brown, weathering blackish grey. Vegetative buds precipitation, some of it as snow, is probably abun-
broadly ovoid; terminal bud 6–10 mm long; lateral dant. On Cerro Potosí, the species forms extensive
buds smaller, slightly resinous. Fascicle sheaths ini- monocultures of close-packed individuals. It occurs
tially 6–8 mm long, in mature fascicles the scales there with scattered, stunted P. hartwegii, which in-
separate and recoil, forming a rosette at the base dicates that the climatic tree line is not reached there
of fascicle, straw-coloured to grey, semi-persistent at around 3700 m. Somewhat lower, on the Sierra La
but mainly falling before the leaf fascicles. Leaves Marta, Coahuila, P. culminicola has been found in
in fascicles of 5 (very rarely 4–6), erect to assurgent, a scrub-community with Quercus spp., Arctostaphy­
persisting 2–3 years, curved, rigid, 3–5 cm long, los, Ceanothus, Agave and grasses; on the Cerro La
0.9–1.3 mm wide, very remotely serrulate to entire, Viega and the Sierra de Arteaga, Coahuila, a similar
obtuse, greyish green, whitish or glaucous adaxially. vegetation, but also with Abies and Pseudotsuga, are
Stomata restricted to both adaxial faces, with 4–5 reported growing scattered with P. culminicula. Pol-
lines on each face. Pollen cones ovoid-oblong when len dispersal has been reported on Cerro Potosí to
mature, 5–8 mm long, yellowish, turning yellowish occur in late July, at 3690 m, which indicates a late
brown. Seed cones subterminal, solitary or in pairs fertilization and short growing season.
on short, stout peduncles which are covered with
subulate-caudate cataphylls. Mature cones subglo-
Conservation
bose when still closed, opening with a flattened base
and remote, spreading fertile seed scales, then 3–4.5 This species is vulnerable to fire during long dry
× 3–5 cm. Seed scales parting and spreading wide periods. In recent years devastating fires have de-
except the smaller, infertile proximal scales, irregu- stroyed large parts of the population on Cerro Po-
lar, often curved, thin, concavo-convex, with 1–2 tosí and regeneration, if it occurs, will be very slow.
deep, cup-like depressions holding the seeds. Apo- IUCN: EN (B1c)
physis slightly raised, transversely keeled, rhomboid
to pentagonal in outline, radially rugose when dry;
Uses
margins crenate or angular, colour yellowish brown,
often resinous. Umbo dorsal, slightly raised, rhom- This species is not used commercially, although lo-
bic in outline, with or without a minute prickle. cally it may be used for firewood. It should be suitable
Seeds obliquely obovoid, 5–7 × 4–5 mm, with a 1 as a low bushy pine for large rockeries, has attractive
mm thick integument, brown. Seed wings absent on foliage, and deserves to be taken into cultivation for
seeds parted from the scales. horticulture. It has established itself well in the rock
garden of the Royal Botanic Garden in Edinburgh,
Scotland, from seed collected by Michael Frankis in
Distribution
1991 and another, slightly later gathering. It remains
Mexico: Coahuila, Nuevo León. very rare in cultivation; it seems that the trade is of-
TDWG codes: 79 MXE-CO MXE-NL ten very slow in recognizing potentially good, albeit
perhaps slow growing, conifers for the garden.
Pinus dalatensis Ferré, Bull. Soc. Hist. Nat.
Taxonomic notes
Toulouse 95: 178. 1960. Pl. 28
Pinus dalatensis is considered by Vietnamese bota-
nist to be a variable species (Nguyen Tien Hiep et
Etymology
al., 2004). This species was studied in the field by
This species is named after Dalat in Vietnam, a city Businský (1999, 2004), who not only discovered sev-
near which it was discovered. eral new populations in formerly unknown locali-
ties, but described them under two subspecies, one
with a new variety. The discriminating characters
Vernacular names
given in his papers are largely continuous and over-
Dalat pine; Thong Da Lat (Vietnamese). lapping, but it seems probable that some of the vari- 659
ation is sufficiently consistent within the disjunct
populations to merit recognition as varieties. I have
Description
retained them here, more or less provisionally, un-
Trees to 30(–40) m tall; trunk to 2(–2.5) m d.b.h., der the ranking given by Businský, and listed only
straight and columnar. Bark smooth in young those few chracter states that appear to more or less
trees and on branches, becoming rough and scaly separate them. It has to be emphasized (again) that
on trunk, breaking into small, grey-brown plates. geographical disjunction is not a taxonomic charac-
Branches wide spreading, self-pruning, forming ter. Some genetic research seems appropriate here.
domed or umbrella-shaped, broad crowns in older
trees. Foliage branches slender, smooth, new shoots
Distribution
glabrous or pubescent, often glaucous or pruinose,
becoming pale brown or reddish brown. Buds small, Vietnam: Binh Tri Thiên, Dac Lac, Gia Lai-Công
conical or ovoid, acute, not resinous; cataphylls ap- Tum, Lâm Dông; Laos: Nakai Nam Theun NPA.
pressed, orange-brown. Leaves in fascicles of 5, with TDWG codes: 41 LAO VIE
deciduous basal sheaths, straight, spreading, (3–)5–
10(–14) cm long, slender, lax or pliant, 0.6–1(–1.2)
Ecology
mm wide; margins mostly minutely serrulate; leaf
colour green or glaucous green (often a combination, Pinus dalatensis is a montane pine occurring at al-
with abaxial face green); apex acute; stomata in fine titudes between 1400 m and 2300 m a.s.l. in tropi-
lines on the two adaxial faces. Pollen cones in small cal evergreen forest. This species forms small stands
clusters at base of new shoots, spirally arranged, of a few to about 30 trees surrounded by evergreen
ovoid to short cylindrical, yellow. Seed cones soli- angiosperm forest, dominated by members of the
tary or in whorls of 2–3 on stout, 1–4 cm long, scaly Fagaceae and Lauraceae. In most localities, the pines
peduncles, persisting some time after seed dispersal, occupy rocky outcrops or steep ridges and adjacent
initially erect but pendulous when grown to full size, slopes where competition from broad-leaved trees is
extremely variable in size, 6–23 cm long, straight or less intense. In more favourable sites on flat moun-
curved ellipsoidal or cylindrical, opening to 5–9 cm tain ridges or in foothills near streams, often on yel-
wide, resinous, green ripening to dull or lustrous low ferralitic soils, it becomes a large, emergent tree.
brown. Seed scales few or numerous depending on Associated conifers found are Pinus krempfii, also an
cone size, flat or boat-shaped, thin woody, flexible; emergent, Fokienia hodginsii, and Dacrydium ela­
basal scales appressed, rarely some recurved. Apo- tum in the southernmost part of the Central High-
physis large, thin or slightly thickened, more or less lands. In most cases, this pine can grow rapidly up
rhombic or with rounded distal margin, straight or to reach the canopy, so it is semi-shade tolerant, but
slightly incurved, striated or grooved towards a ter- seedlings are intolerant of shade, so some form of
minal, 4–10 mm wide, sunken, flat or obtuse umbo. canopy disturbance has to precede regeneration, e.g.
Seeds varying in size with the seed scales (especially periodic fires.
the wings), with maximal lengths of 8–10 mm and
wings to 20–30 mm long and 10 mm wide, the latter Conservation
often with dark longitudinal streaks.
This species has recently been conservation assessed
(Nguyen Tien Hiep et al., 2004) and is considered to
660

Pl ate 2 8 . Pinus dalatensis. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cones, one green,
one ripe. 5. Seeds.
be Vulnerable. Populations are few and small, and
Distribution
several have been in continuous decline; on the oth-
er hand, given recent discoveries and their disjunct Vietnam: Lâm Dông (Bi Doup Mountain).
occurrence, new populations are likely to be found. TDWG codes: 41 VIE
Some of these are in as yet undisturbed forests. The
main threat in the past was logging, often as an ad-
Conservation
junct to the much sought after large trees of Fokienia
hodginsii; today it is mostly conversion to managed IUCN: DD
pine forest with planted Pinus kesiya, or conversion
of primary forest to other land uses via slash-and- Pinus dalatensis Ferré subsp. procera Businský,
burn agriculture. Several populations are within re- Candollea 54: 133. 1999. Type: Vietnam: Gia Lai 661
cently declared national parks and nature reserves. Prov., Ngoc Linh Mountains, Mt. Ngok Niay,
IUCN: VU (B1+2c) R. Businský 44114 (holotype G).

Uses Description
A valuable timber tree, but very rare and logged only New shoots (mostly densely) pubescent. Seed cones
incidentally. Its wood properties are similar to those 9–23 cm long.
of Pinus wallichiana, a native of the Himalayas, and
other so-called white pines. It is not known in cul-
Distribution
tivation.
Vietnam: Gia Lai-Công Tum (Ngoc Linh, Ngok Niay
2 subspecies and 2 varieties are recognized: Mountain, Kon Plong); perhaps across the border in
Laos.
Pinus dalatensis Ferré subsp. dalatensis var. dalat­ TDWG codes: 41 VIE
ensis. Type: Vietnam: Lâm Dông, Da Lat, Trai Mat,
H. Gaussen s.n. (holotype TLF).
Conservation
Description IUCN: DD
Note on conservation status: Although the species
New shoots variably pubescent. Seed cones 6–16 cm
as a whole has been assessed as VU, the status of the
long.
separate infraspecific taxa, although logically meet-
ing the criteria for at least VU, remains at present
Distribution
unknown because of insufficient data.
Vietnam: Binh Tri Thiên, Dac Lac, Lâm Dông (Chu
Yang Sinh Massif, Da Lat environs); Laos: Nakai
Nam Theun NPA. Pinus densata Mast., J. Linn. Soc., Bot. 37: 416.
TDWG codes: 41 LAO VIE 1906. Pinus tabuliformis Carrière var. densata
(Mast.) Rehd., J. Arnold Arbor. 7: 23. 1926. Type:
Conservation China: Sichuan, Yalong River, E. H. Wilson 3015
(holotype BM).
IUCN: DD

Pinus dalatensis Ferré subsp. dalatensis var. bi­ Etymology

doupensis Businský, Candollea 54: 127. 1999. Type:
The species epithet is perhaps from Latin densatio =
Vietnam: Lâm Dông, Dong Nai, Mt. Bi Doup,
thickening (condensation) and may refer to the apo-
W slopes of massif 6 km SE of Bi Doup, R. Businský
physes; if so it is not used in its original sense.
39160 (holotype G).

Description Vernacular names

New shoots glabrous. Seed cones 10–16 cm long. Gaoshan pine; gao shan song (Chinese)
 in plants generally, we would not gain much relevant
Description
information if we did prove it.
Trees to 30 m tall; trunk to 1.3 m d.b.h. Bark scaly,
forming irregular plates and longitudinal fissures,
Distribution
dark grey-brown. Branches spreading, forming a
broadly domed or open crown in older trees. Foli- China: S Qinghai, W Sichuan, NW Yunnan, E Xi-
age branches stout, new shoots glabrous, rough with zang [Tibet].
pulvini from fallen leaf fascicles, lustrous yellowish TDWG codes: 36 CHC-SC CHC-YN CHQ CHT
brown, turning reddish brown. Buds ovoid-conical,
acute, the terminal bud ca. 15 mm long, slightly res-
Ecology
662 inous; cataphylls with white fringed margins, acumi-
nate. Leaves in fascicles of 2(–3), held by a persistent, Pinus densata occurs in high mountains at altitudes
10–15 mm long but shortening basal sheath, straight, from 2600 m to 4000 m a.s.l. or even above this line
8–14 cm long, rigid, slightly twisted, 1–1.5 mm wide, to 4200 m. It forms open pure stands at the highest
dull green; margins minutely serrulate; apex acute; elevations, but becomes mixed with other pines, e.g.
stomata in fine lines on all faces. Pollen cones in P. armandii and P. yunnanensis, below 3000 m a.s.l.
clusters at base of new shoots, spirally arranged, It is presumed that its hybrid characters include a
short cylindrical, ca. 2 cm long, yellow. Seed cones greater tolerance to frost than either of its putative
solitary or in pairs, sessile or very short pedunculate, parents, P. tabuliformis and P. yunnanensis, enabling
persistent but opening widely, spreading or curved it to colonize right up to the present tree line in
down, narrowly ovoid when closed, 4–6 cm long, western China’s great mountain system.
when opened 4–7 cm wide. Seed scales oblong, thin
woody, rigid, brown; apophyses prominently raised,
Conservation
more or less rhombic, strongly transversely keeled,
lustrous chocolate brown; umbo dorsal and central, IUCN: LC
armed with a short prickle. Seeds ovoid to ellipsoid,
4–6 mm long, light brown to grey-brown; wing 15–
Uses
20 mm long.
There is presently little economic use of this species,
primarily because it occurs in high mountains in
Taxonomic notes
often inaccessible places. A traditional use at least
Maxwell Masters (op. cit.) described this pine as a century ago seems to have been making torches,
a new species and compared it with Pinus den­ using wood and resin. It has been suggested that
siflora and to some extent (via E. H. Wilson’s field it would be suitable for afforestation (presumably
observations) with P. sylvestris. Both species occur forest restoration) in the alpine regions of western
far to the NE from the known range of P. densata. China. It was introduced in Europe (UK) about 100
The hybrid status of this species has later been in- years ago, but this species remains rare in botanic
ferred from morphological intermediacy of charac- gardens and arboreta.
ter states in comparison with P. yunnanensis, with
which it is partially sympatric, and P. tabuliformis,
which occurs also further north in China. Some Pinus ×densithunbergii Uyeki, [Not. Pl. Lign.
molecular evidence (DNA sequence data) supports Sikoku 2] Sci. Rep. Matsuyama Agric. Coll. 10: 6.
this hypothesis, possibly with the involvement of an 1953. Pinus densi-thunbergii Uyeki, Corean Timber
unknown, third species from the Tertiary. Due to Trees 1: 65 (Chôsen 4). 1926. Type: Illustration in
ecological as well as geographical separation, there Uyeki, Corean Timber Trees 1: photograph opp.
is no indication that these species form new hybrids p. 66, No. 1–6. 1926 (lectotype designated here).
at present in their natural habitats. It seems to me
that it is appropriate to treat this taxon not as a not-
Etymology
hospecies but as a species. Its hybrid status has not
been demonstrated beyond doubt, and given that The nothospecific epithet combines the names of
hybridization is probably a major cause of ­speciation the parental species.
 ­ inus densiflora Siebold & Zucc. var. sylvestriformis
P
Vernacular names
(Taken.) Q. L. Wang, Changbai Shan zhi-wu min-lu
Aiguro-matsu, Akakuro-matsu (Japanese) [Checklist of plants in Changbai Mt.]: 49. 1982.
Pinus densiflora Siebold & Zucc. var. zhangwuensis
S. J. Zhang et al., Bull. Bot. Res. North-East. Forest.
Description
Inst. 15 (3): 338. 1995.
A natural natural hybrid between Pinus densiflora
Siebold & Zucc. and P. thunbergii Parl, first described
Etymology
by Mayr (1890) and named first as a species and then
as a nothospecies by Uyeki. Its morphological char- The species epithet means ‘with compact inflores-
acters are described as intermediate between the cence’ and may refer to the closely set pollen cones. 663
two species. The resin ducts are external (marginal)
in Pinus densiflora and medial in P. thunbergii; in
Vernacular names
the leaves of the hybrid taxon both positions were
found. The thickness of the leaf hypodermis is de- Japanese red pine; Akamatsu (Japanese); chisong
scribed as intermediate between those of the parent (Chinese); sonamoo (Korean)
species.
Description
Distribution
Trees to 30–35 m tall, in coastal areas smaller, to
Japan: Honshu, Shikoku, Kyushu; South Korea. 20 m tall; trunk to 1–1.5 m d.b.h., bole straight and
TDWG codes: 38 JAP-HN JAP-SH JAP-KY KOR-SK columnar, self-pruning. Bark of young trees and in
crown of old trees thin, with papery flakes, orange-
red; on large trunks becoming thick and breaking
Ecology
into large, irregular plates, turning grey-brown
Apparently widespread among populations of both or red-brown. Branches spreading, higher order
putative parents. branches assurgent, forming an umbrella-shaped or
flat-topped crown. Foliage branches slender, young
shoots smooth, glabrous, yellowish, sometimes
Conservation
slightly glaucous, becoming rough with persistent
IUCN: LC pulvini from fallen leaf fascicles, turning dark grey-
brown to grey. Buds ovoid-oblong, slightly resinous;
cataphylls dark red-brown. Leaves in fascicles of 2,
Uses
persisting 2–3 years, held by a short basal sheath,
Controlled crossings of Pinus densiflora with P. thun­ straight and rigid, 6–12(–15) cm long, often slightly
bergii have been undertaken in Japan and the USA. twisted, ca. 1 mm wide; margins minutely serrulate;
apex acute-acuminate; leaf colour green or glaucous
green; stomata in fine lines on all surfaces. Pollen
Pinus densiflora Siebold & Zucc., Fl. Japon. 2 (3): cones in small clusters at base of new shoots, spirally
22, t. 112. 1842. Type: Japan: [“in Japonia”], P. F. von arranged, short cylindrical, 1.5–2.5 cm long, yellow.
Siebold s.n. comm. 1842 ex herb. Zuccarini No. 438 Seed cones mostly solitary, erect or reflexed on short
(lectotype M). peduncles or nearly sessile, falling after seed disper-
sal, symmetrical or nearly so, 3–6 cm long, ovoid-
Pinus funebris Kom., Trudy Imp. S.-Peterburgsk. conical when closed, broadly ovoid when opened.
Bot. Sada 20: 177. 1901; Pinus densiflora Siebold & Seed scales thin woody, rigid, oblong, spreading
Zucc. var. funebris (Kom.) T. N. Liou & Q. L. Wang moderately wide; apophyses flat or slightly raised,
ex Silba, Phytologia Mem. 7: 50. 1984. transversely keeled, more or less rhombic in outline
Pinus densiflora Siebold & Zucc. f. sylvestriformis or with upper margin rounded, dull brown; umbo
Taken., J. Japan. Forest. Soc. 24: 120, f. 1. 1942; Pinus broadly triangular or irregular, depressed or slightly
sylvestris L. var. sylvestriformis (Taken.) W. C. Cheng protruding, unarmed or with a weak, minute prick-
& C. D. Chu, Fl. Reipubl. Pop. Sin. 7: 246. 1978; le. Seeds obovoid or ellipsoid, slightly flattened, 4–7
 mm long, dull brown or tan; wing firmly attached,
Uses
12–20 mm long, 5–7 mm wide.
Japanese red pine is very similar (and closely relat-
ed) to Scots pine and has consequently similar wood
Taxonomic notes
properties; it is an important timber tree in NE Asia.
Pinus funebris has been described by Komarov (op. The wood is today mainly used in the paper indus-
cit.) as a separate species from North Korea and the try, but also still provides timbers for underground
adjacent Russian border area; some later authors mining and for railway sleepers, as well as construc-
considered it to be a variety of P. densiflora. Quite a tion timber. Foresters in Japan and the USA have
number of other forms or varieties were subsequent- produced hybrids with P. thunbergii and P. massoni­
664 ly described and named, all from NE China. None ana and with P. nigra as well as with P. sylvestris. In
of these are recognized here, but Flora of China 4 Japan, this species is extensively planted for forestry
(1999) still recognizes two; one of these, P. densi­flora as well as for amenity; in Japanese horticulture per-
var. ussuriensis T. N. Liou & Q. L. Wang is a synonym haps as many as 100 cultivars are known. This pine
of P. sylvestris (Farjon, 1998, [2001]). The other varie- and its cultivars are often used in Japanese landscape
ty, var. zhangwuensis S. J. Zhang et al., lacking “white gardens of larger size, traditionally around shrines
powdery” (glaucous?) branchlets, is described with and in palace grounds. Relatively few of these culti-
longer (13–15 cm) needles than var. densiflora (6–12 vars, and indeed the species itself, have made their
cm), but unless (and until) such descriptions are way to Europe, probably because there P. sylvestris
based on wide sampling, enabling morphometric offers similar opportunities for horticultural experi-
studies, they remain unconvincing. mentation. The Japanese horticultural art ‘bonsai’
also makes use of this species. Needles and extracts
from them are used in traditional medicine such as
Distribution
aromatherapy. The pollen is edible and also used as
China: Heilongjiang, Jiling; Japan: Honshu, Kyushu, medicine.
Shikoku; North & South Korea; Russian Far East:
Primorye.
TDWG codes: 31 PRM 36 CHM-HJ CHM-JL 38 JAP- Pinus devoniana Lindl., Edwards’s Bot. Reg. 25:
HN JAP-KY JAP-SH KOR-NK KOR-SK 62. Aug 1839. [Allg. Gartenzeitung 7: 324. 1839]
Type: Mexico: Hidalgo, Mineral del Monte, Cerro
Ocotillo, C. T. Hartweg s.n. (lectotype W). Fig. 222
Ecology
Pinus densiflora occurs in extensive pure stands in Pinus michoacana Martínez, Anales Inst. Biol. Univ.
many parts of its range and is one of the most domi- Nac. México 15: 1. 1944.
nant conifers in Japan and Korea. It grows in a vari-
ety of acidic soils, from dry sandy or rocky sites to
Etymology
peaty soils. In Japan it reaches from near sea level
(and close to the shore) up to 2300 m in the moun- This species was named in honour of William Spen-
tains, but on mainland Asia its altitudinal range is cer Cavendish, sixth Duke of Devonshire (1790–
more restricted and extends upward to only 900 1858), who had a great interest in gardening.
m a.s.l. in NE China and 1300 m in Korea. In areas
where broad-leaved forest dominates, P. densiflora is
Vernacular names
restricted to poorer sites such as rock outcrops on S-
facing slopes and edges of moors or mountain lakes. Michoacan pine; pino blanco, pino lacio, ocote gre-
Here it mixes with the angiosperms and can quickly tado (Mexico)
recolonise ahead of them after forest fires.
Description
Conservation
Trees to 20–30 m tall, d.b.h. to 0.8–1 m. Trunk mo-
IUCN: LC nopodial, usually erect; bark thick, scaly, very rough,
with elongated plates divided by deep longitudinal
black fissures, reddish brown to dark brown. Branch- available name Pinus devoniana Lindley. Hopefully,
es spreading and assurging, or ascending near the the results of thorough taxonomic work published
top, forming an open, broad pyramidal or domed in specialized literature that strictly follows the
crown. Shoots 15–20 mm thick, rigid, curved, very rules of botanical nomenclature will eventually filter
rough and scaly. Cataphylls up to 20–25 mm long, through to become established in the literature aim-
subulate, scarious, recurved or reflexed, dark brown ing at a more general audience.
to blackish grey, with erose-ciliate margins. Vegeta-
tive buds large; terminal bud 20–40 mm long, but
Distribution
lateral buds smaller, ovoid-acute, not resinous. Fas-
cicle sheaths to 30–40 mm long, often very resin- Mexico: Sinaloa, Nayarit, Jalisco, Zacatecas, Agua­
ous. Leaves in fascicles of 5, rarely 4 or 6, persisting scalientes, San Luís Potosí, Querétaro, Hidalgo, 665
2–3 years, rigid, straight, or flexible and drooping, Michoacán, México, Distrito Federal, Morelos,
(17–)25–40(–45) cm long, 1.1–1.6 mm wide, with Tlaxcala, Puebla, Veracruz, Guerrero, Oaxaca and
serrulate margins, acute-pungent, lustrous green. Chiapas; in Guatemala in the southern highlands.
Stomata on all faces of leaves. Pollen cones cylindri- TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
cal, 20–40 mm long, pink-purplish, turning light PU MXC-TL MXE-AG MXE-HI MXE-QU MXE-SL
brown. Seed cones subterminal, solitary or in pairs MXE-ZA MXG-VC MXN-SI MXS-GR MXS-JA MXS-MI
or whorls of 3–4. Mature cones on short, persistent MXS-NA MXS-OA MXT-CI 80 GUA
peduncles, leaving a few scales on the branch when
falling, variable in size and shape, typically ovoid-
Ecology
oblong, with an oblique base, or curvate to cornute
(often by insect-damage), 15–35 × 8–15 cm when This species is a constituent of relatively open, of-
open. Seed scales parting when the seeds are ma- ten secondary pine-oak forests, or it occurs with
ture, spreading wide, straight or slightly recurved, Pinus oocarpa invading burned mountainsides.
thick woody, rigid or flexible with some force. Apo- It is well adapted to withstand fires by its juvenile
physes mostly raised, transversely keeled, rhombic “grass stage” similar to that of P. palustris of the SE
in outline, with irregular sides, up to 25 mm wide, USA. The pines most commonly associated with
often radially striated, in various shades of brown. it vary somewhat from north to south and include
Umbo dorsal, raised, flat or depressed, terminating P. oocarpa, P. montezumae, P. pseudostrobus, and
in a small, usually deciduous prickle, grey-brown. P. maximinoi and less frequently P. cembroides at
Seeds obliquely broad ovoid, flattened, 8–10 × 5–7 lower altitudes and P. hartwegii at higher altitudes.
mm, light brown, often with dark spots. Seed wings Common are also Quercus, Liquidambar, and in the
obliquely ovate to oblong, with a straight side, 25–35 understorey Calliandra, Leucaena, Acacia, Dodo­
× 10–15 mm, light brown with darker stripes. Many naea, Gaultheria, and Mimosa. Its altitudinal range
seedlings developing a ‘grass stage’ with delayed api- is (700–)900–2500(–3000) m a.s.l. Pinus devoniana
cal growth of stem. grows on a variety of soils, often of volcanic origin.
The climate is warm-temperate to subtropical, with
annual precipitation 1000–1500 mm and a dry sea-
Taxonomic notes
son from November to May.
This species is still known in many books on coni-
fers by its name given by Maximino Martínez, Pi­
Conservation
nus michoacana. Martínez (1945, 1948), who worked
on the pines of Mexico during the difficult years of IUCN: LC
World War II, considered it likely that his new spe-
cies would correspond with some of John Lindley’s
Uses
names, among which P. devoniana was mentioned,
but at the time he had no access to these original Pinus devoniana is a common tree throughout the
publications. The complexity of forms and many of mountainous parts of S Mexico and Guatemala, of-
the names that were given to these have been united ten encountered in open, degraded pine-oak or pine
by Farjon & Styles (1997) in a comprehensive mono- forest. As it is not a very tall, straight-boled tree and
graph on Latin American pines under the earliest has branches low on the trunk, it is not often ­selected
 as a timber tree. On the other hand, it is locally used faces of leaves. Pollen cones numerous, forming
and, like other pines, increasingly for firewood by long, spiculate clusters, cylindrical, 20–25 × 4–5
the growing rural population. Uses of timber are mm, pinkish brown at maturity. Seed cones solitary,
put to fence posts, boxes, furniture, toll handles, and in pairs or whorls of 3–4 on stout, recurved, 10–15
other woodware and, mixed with other pine wood, mm long peduncles which fall with the cone. Mature
wood chips glued and compressed to particleboard. cones ovoid to ovoid-attenuate when closed, often
It is virtually unknown in horticulture despite the slightly curved, more broadly ovoid with a flattened
fact that it is one of the most strikingly beautiful base when opened, then 7–10 × 5–7 cm. Seed scales
pines, with perhaps the longest needles in the entire parting to release the seeds except at base, (thin)
genus and large cones and a deep red-brown bark. woody, oblong, straight or slightly curved. Apo­
666 Provenance is crucial to establish whether some physes nearly flat or raised and transversely keeled,
trees could turn out to be hardy, but otherwise it radially striate or grooved, rhombic or pentagonal in
would do fine in much of California, the Mediter- outline, nearly symmetrical around cone, 11–17 mm
ranean, Australia, and New Zealand, to mention just wide, light brown, reddish brown or dark brown.
a few regions where gardening and tree planting are Umbo dorsal, raised, transversely rhombic, darker
a major enjoyment for many. than the apophysis, prickle absent. Seeds obliquely
obovoid, slightly flattened, 4–5 × 3–3.5 mm, light
grey or brown, often with dark spots. Seed wings ob-
Pinus douglasiana Martínez, Madroño 7: 4. 1943. long, with a straight and a curved side, 18–24 × 7–9
Type: Mexico: Nayarit, Jalá, Cerro Juanacata, mm, light yellowish brown, translucent, sometimes
M. Martínez 3429 (holotype MEXU). with a darker brown tinge.

Etymology Distribution
This species was named after the plant collector Mexico: mainly in Jalisco, Michoacán, México and
David Douglas (1798–1834), who, however, never N Morelos, but extending northward into Nayarit
visited Mexico. and the crest of the Sierra Madre Occidental on the
border line of Sinaloa and Durango, southwards it
occurs locally in Guerrero and Oaxaca.
Vernacular names
TDWG codes: 79 MXC-ME MXC-MO MXC-PU MXE-
Douglas pine; ocote, pino hayarín (Mexico) DU MXN-SI MXS-GR MXS-JA MXS-MI MXS-NA MXS-
OA
Description
Ecology
Trees to 20–45 m tall, d.b.h. to 0.8–1 m. Trunk
mono­podial, straight; bark rough, scaly, divided This species is a constituent of mostly mixed pine
into large, irregular plates and deep fissures, dark or pine-oak forests at elevations ranging from
reddish brown, weathering grey-brown. Branches (1100–)1400–2500(–2700) m a.s.l. in warm to tem-
long, slender, spreading, or ascending in upper part perate climatic zones. The annual precipitation dif-
of crown. Shoots uni-nodal, dark brown, not glau- fers with altitude but is roughly around 1000 mm
cous. Cataphylls subulate-caudate, spreading or in most areas. Common associated pines are Pinus
recurved, with erose-ciliate margins, brown. Vegeta- pseudostrobus, P. herrerae, P. leiophylla, P. lawsonii,
tive buds ovoid-conical; terminal bud 15–25 × 10–15 P. ayacahuite in the southern part of its range, some-
mm, but lateral buds smaller, not resinous. Fascicle times P. oocarpa and P. devoniana at lower elevation
sheaths persistent, remaining long, (15–)20–35 mm, and drier sites respectively. At the highest and wet-
weathering greyish brown. Leaves in fascicles of 5, test sites P. douglasiana can occur with Abies, Picea
rarely 4 or 6, persisting 2–2.5 years, slender, lax or (in Durango) or Cupressus lusitanica. In many areas
sometimes more rigid, 22–35 cm long, 0.7–1.2 mm Quercus spp. are codominant; a shift in forest com-
wide; margins serrulate; apex acute; leaf colour light position towards broad-leaved trees may occur also
yellowish green to glaucous green. Stomata on all by selective cutting of pines.
 persistent but reduced to 10–15 mm. Leaves in fas-
Conservation
cicles of (4–)5–6(–7, rarely 8), persisting 2–2.5(–3?)
IUCN: LC years, straight or slightly curved, 14–24 cm long,
0.7–1.1 mm wide; margins serrulate, acute to pun-
gent; leaf colour yellowish green to glaucous green.
Uses
Stomata on all faces of leaves. Pollen cones ovoid-
Pinus douglasiana is, along with other species with oblong to cylindrical, yellowish brown when ma-
which it often occurs, an important timber tree ture. Seed cones subterminal, solitary, in pairs or in
in most of its range. It is, however, not especially whorls of 3–4 on short, stout, usually persistent pe-
sought out for tree-breeding purposes, perhaps due duncles, eventually deciduous. Mature cones ovoid
to the fact that it is relatively unknown to foresters. or broadly ovoid when opened, slightly flattened 667
It is perhaps present in a few arboreta or pineta in or obtuse-conical at base, often slightly curved,
regions with a mild climate, but not generally cul- 5–9(–11) × 4–6(–7) cm when open. Seed scales part-
tivated for horticulture. It has been tried in forestry ing readily to release seeds except those at base of
plantations, but is not in common use. cone, thick woody, broadly oblong, straight or re-
curved near base of cone. Apophysis raised, some-
times flat on basal scales, prominently transversely
Pinus durangensis Martínez, Anales Inst. Biol. keeled, rhombic to pentagonal in outline, more or
Univ. Nac. México 13: 23, f. 1–4. 1942. Type: Mexico: less symmetrical around cone, ochraceous to light
Durango, El Salto, Las Adjuntas, M. Martínez 3477 (reddish) brown. Umbo dorsal, raised, slightly re-
(holotype MEXU). Fig. 223 curved, transversely keeled, with a small, persist-
ent (or deciduous) prickle. Seeds obliquely ovoid,
Pinus martinezii E. Larsen, Madroño 17: 217. 1964; slightly flattened, 5–6 × 4–4.5 mm, light brown to
­Pinus douglasiana Martínez var. martinezii (E. Lar­ grey, with small, dark spots. Seed wings oblique,
sen) Silba, Phytologia 68: 49. 1990. 14–20 × 6–9 mm, light greyish brown, translucent,
with a blackish tinge.
Etymology
Distribution
The species epithet durangensis refers to the Mexican
State of Durango, from where it was first described. Mainly in the southern Sierra Madre Occidental of
Mexico, rare in E Sonora and Chihuahua, common
in Durango, Zacatecas and N Jalisco, more scattered
Vernacular names
further south in Jalisco and N Michoacán.
Durango pine; ocote, pino blanco, pino real (Mex- TDWG codes: 79 MXE-CU MXE-DU MXE-ZA MXN-
ico) SI MXN-SO MXS-JA MXS-MI

Description Ecology
Trees to 35–40 m tall, d.b.h. to 0.8–1 m; trunk mo- In the Sierra Madre this species is an important con-
nopodial, erect, straight. Bark rough, scaly, breaking stituent of the ‘yellow pine’ forest, where it occurs
into large, irregular, elongated plates divided by lon- in pure stands or mixed with several other species
gitudinal, shallow fissures, dark brown, weathering of pine, e.g. P. arizonica, P. leiophylla, and P. engel­
grey. Branches long, slender, with lower branches mannii, or in pine-oak forests. Its altitudinal range
usually curved downward, ultimately more or less is extensive: (1400–)1600–2800(–3000?) m, but
pendant. Shoots uni-nodal, orange-brown or red- it is most common at 2000–2800 m a.s.l. Climati-
dish brown, usually glaucous. Cataphylls ca. 15 mm cally, the Sierra Madre is warm-temperate, but with
long, 3–4 mm wide at base, subulate, soon reflexed, cold spells during the short winter at the higher el-
scarious, with erose-ciliate margins and caudate evations. Annual precipitation varies between 700–
apex, dark brown. Vegetative buds ovoid; terminal 1200 mm, most of which occurs in the summer. This
bud 15–20(–25) × 10–15 mm, but lateral buds small- pine is adapted to grow on shallow, rocky soils, but
er, not resinous. Fascicle sheaths 20–30 mm long, its better stands are found on deeper soils, where it
 can successfully compete with most other pines. The spreading, sometimes heavy but more slender and
soils are mostly derived from volcanic rock. At the shorter in trees growing in closed canopy stands. Fo-
highest elevation P. durangensis occurs with Abies liage branches slender, often more or less pendulous,
and/or Cupressus lusitanica, at the lowest with Juni­ new shoots purplish green, often glaucous, turning
perus deppeana and Pinus oocarpa. Other pines are red-brown to grey, rough with persistent pulvini
P. montezumae, P. teocote and in the southern part of after leaf fascicles have fallen. Buds ovoid to ovoid-
its range P. ayacahuite can occur with it. cylindric, 5–10 mm long, strongly resinous. Leaves
in fascicles of 2 or 3, held in a 10–15 mm long fascicle
sheath, persisting 3–5 years, spreading, (5–)6–11(–12)
Conservation
cm long, straight or slightly curved and flexible, ca.
668 Pinus durangensis, as a pine tree with a tall, straight 1 mm wide, yellowish green, greyish green or dark
bole, has been much sought after as a timber tree green; margins minutely serrulate; apex short acute;
and exploitation has been severe, especially in Du- stomata in fine lines on all surfaces. Pollen cones in
rango (Perry, 1991), where extensive, more or less small clusters, spirally arranged, cylindrical, 1.5–2
pure stands are now rare. In mixed forest and remote cm long, pale purplish green to yellow, turning light
areas mature to aged trees still occur abundantly, but brown. Seed cones solitary or in whorls of 2–4, short
in the near future protection may become necessary pedunculate or nearly sessile, semi-persistent but
to avoid over-exploitation. opening soon, spreading, 4–7 cm long, narrowly
IUCN: LC ovoid or ovoid-oblong when closed, broadly ovoid
or ovoid-conical with a nearly flat base when open.
Seed scales thin woody, rigid, uniformily dull brown
Uses
below the apophysis without a dark band; apophysis
Durango pine is an important timber tree. It grows slightly raised, sharply keeled transversally; umbo
straight and tall and is (or was) abundant and wide- dorsal and central, armed with a short, stout prickle.
spread in many areas within its range. Logging from Seeds ellipsoid, ca. 6 mm long, grey mottled black,
natural stands at the current rate is unsustainable with a similarly patterned wing 12–15 mm long.
and plantations are now being attempted in the state
of Durango. The timber is used for construction
Distribution
such as roof beams, general carpentry, furniture,
floors, and plywood. This species is virtually un- E and SE USA, from New York to E Texas accross
known in horticulture. 22 states.
TDWG codes: 74 MSO OKL 75 CNT NWJ NWY OHI
PEN WVA 77 TEX 78 ALA ARK DEL FLA GEO KTY
Pinus echinata Mill., Gard. Dict., ed. 8: Pinus No. LOU MRY MSI NCA SCA TEN VRG WDC
12. 1768. Type not designated.
Ecology
Etymology
Pinus echinata is a lowland pine with an extensive
The species epithet means ‘spiny’ or ‘prickly’, but it range across the SE United States, mostly growing
is not clear what Philip Miller found so spiny in this from ca. 150 m up to the foothills of the Appalachian
pine. Mountains at ca. 600 m a.s.l. It is absent in the Mis-
sissippi Valley and its delta as well as in a narrow
to fairly wide coastal strip along the Gulf of Mexico
Vernacular names
and Atlantic Ocean, and it does not extend into most
Shortleaf pine of Florida. This indicates that it is primarily limited
by climatic factors, such as the 10 °C average annual
temperature isoline at its northern limit and an aver-
Description
age annual precipitation above 1000 mm distributed
Trees to 35–40 m tall; trunk to 1.6 m d.b.h. Bark more or less evenly over a year as its southern limit.
breaking into irregularly rectangular or square, It grows on a great variety of soils, but most have a
scaly plates, dark brown, slightly resinous. Branches capacity for moist retention with a sandy loam or
silty loam texture and good drainage. Although this
Description
species can form nearly pure stands, in most sites
it will be succeeded by broadleaved trees especially Shrubs or trees to 15(–21) m tall, d.b.h. to 60 cm.
oaks (Quercus spp.) except where thin soil overlies Trunk monopodial, short to medium size, strongly
rock. There P. echinata forms a minor component tapering, branching low. Bark thick, rough and scaly,
of various angiosperm-dominated forest and wood- breaking into small, irregular plates, divided by ir-
land types. It can also be associated with P. taeda, regular, shallow fissures, not easily exfoliating, red-
which has a very similar distribution, but at least in brown weathering grey-brown. Branches spreading
parts of its range occurs in a wetter habitat. or ascending, irregularly disposed in most trees.
Crown broad conical or rounded, dense. Shoots pale
red-brown or tan, rarely slightly glaucous at first, 669
Conservation
soon grey, glabrous or minutely puberulent. Vegeta-
IUCN: LC tive buds ovoid-oblong to ellipsoid; terminal bud
5–10 × 3–5 mm, but lateral buds smaller, resinous,
red-brown. Fascicle sheaths 5–7 mm long, loosely
Uses
imbricate, soon recoiling, then pale straw-coloured
Shortleaf pine is an important commercial conifer to grey, forming a small rosette at base of fascicle but
species in the SE United States and both natural deciduous before the leaves fall. Leaves in fascicles of
stands and plantations are exploited for timber. The (1–)2(–3), spreading, persisting 4–6 years, upcurved,
wood is of excellent quality, with orange or yellow- connivent, rigid, 2–4 cm long, 0.9–1.5 mm wide;
ish brown heartwood and creamy yellow sapwood; margins entire or minutely serrulate; leaf colour var-
it is used for railway sleepers, construction lumber, iable, dull green to glaucous green. Stomata: leaves
indoor finishing like panelling, plywood, furniture, amphistomatic, with 2–4 lines on each face. Pollen
and kraft pulp and dissolving pulp; the latter product cones in more or less elongated, spiculate clusters,
feeds the paper industry. Most of the plantation tim- ca. 7 × 3.5 mm, yellowish to red-brown. Seed cones
ber goes to pulping. There is a limited use for amen- solitary, paired or more rarely in whorls of 3, on very
ity planting and in urban areas this pine is planted to short, 3–5 mm long peduncles remaining with fallen
screen off residential areas from motorways (major cones. Mature cones seemingly sessile, irregularly
highways) or industrial areas. This species has little globose or ovoid-globose when closed, spreading
significance in horticulture and is rarely planted in often wider than long when opened, with a flattened
gardens. base, irregular in size and shape, often resinous, 3–5
× 3–6 cm when open. Seed scales parting easily and
widely, narrowly and more or less weakly attached
Pinus edulis Engelm., in Wislizenus, Mem. Tour to the rachis and hence moveable, concavo-convex,
N. Mexico: 88. 1848. Pinus cembroides Zucc. var. with 1–2 deep seed cavities. Apophysis prominent-
edulis (Engelm.) Voss, Mitt. Deutsch. Dendrol. Ges. ly raised, transversely keeled or radially keeled or
1907 (16): 95. 1907; Pinus cembroides Zucc. subsp. ribbed, rhombic to pentagonal in outline but irregu-
edulis (Engelm.) E. Murray, Kalmia 12: 22. 1982. lar, often angular or curved, variously brown. Umbo
Type not designated. dorsal, flat or slightly raised, with a minute prickle.
Seeds obovoid or ellipsoid, 10–15 × 6–9 mm, greyish
brown or light brown; integument thick, 0.5–1(–1.1)
Etymology
mm; megagametophyte (‘endosperm’) pinkish or
The species epithet means ‘edible’ and refers to the white when fresh. Seed wings absent when the seed
seeds. is detached from the scale.

Vernacular names Distribution

Pinyon pine, Two-needle pinyon pine, Colorado SW USA: Arizona, S California, Colorado, New
pinyon Mexico, W Oklahoma, NW Texas, Utah, S Wyoming.
TDWG codes: 73 COL WYO 74 OKL 76 ARI CAL
UTA 77 NWM TEX
 burns with a distinctive aroma (as does the juni­
Ecology
per wood, which I prefer for my campfires when
Pinus edulis is widely distributed in the interior ba- traveling in the area). Large quantities were logged
sins, plateaus, mesas and mountains of the ‘Four and its rough timbers used as props in the mining
Corner’ States of the USA. It forms extensive, open boom of the late 19th century. Poor growth form
stands commonly with one or more species of Juni­ from the forester’s perspective renders the wood
perus known as Pinyon-Juniper woodland, which unsuitable for sawn timber, even though its quality
is one of the most widespread semi-arid vegetation matches that of Ponderosa pine. The edible seeds are
types in North America. Summers are hot and win- easy to harvest and in great demand as a delicacy;
ters cold, but climatic conditions are varying with they may constitute the economically most valuable
670 altitude and latitude. Soils are commonly thin to product of the species. Crops can vary greatly from
sceletal or may be absent altogether, with the trees one year to the next and the slow growing pines do
growing from fissures in the sandstone, limestone, not perform well in plantations. Horticultural value
or shale. Recent sedimentation accumulates in the is limited, although locally young trees are harvested
basins and valley bottoms, where grasses and ‘sage- as Christmas trees.
brush’ (Seriphidium tridentatum) dominate, while
at higher elevations in the mountains the Pinyon-
Juniper woodland gives way to open pine forest with Pinus elliottii Engelm., Trans. Acad. Sci. St. Louis 4:
Pinus ponderosa and Pseudotsuga menziesii. Junipe­ 186, t. 1–3. 1880.
rus monosperma and J. osteosperma are the most
commonly associated junipers with P. edulis. The
Etymology
appearance is of a stunted forest as the free standing
trees branch low and form wide spreading crowns This species was named after the botanist Stephen
while only attaining modest height. Depending Elliott (1771–1830).
on openess, there is an understorey dominated by
shrubs of which Seriphidium (Artimisia) is most
Vernacular names
common and widespread, supplemented by scrubby
oaks (Quercus spp.), Chrysothamus, Cercocarpus, Slash pine
Ephedra, Yucca, and several others depending on
geographical area, as well as grasses and other herbs.
Description
Trees to 30 m tall; trunk to 80 cm d.b.h., straight or
Conservation
contorted. Bark breaking into large, irregularly rec-
Although the species is too widespread and abun- tangular, thin papery or scaly plates separated by ir-
dant to be threatened with extinction, the Pinyon- regular furrows, orange-brown or purplish brown.
Juniper woodland as an ecosystem is under threat Branches spreading, forming a wide, dome-shaped,
in many places due to ‘range improvement’ for the open crown. Foliage branches stout, often to ca. 1 cm
grazing of cattle and sheep, causing the removal or thick, orange-brown, turning red-brown to dark
degradation of the woodland over large areas. There brown, rough with persistent pulvini after leaf fas-
is growing awareness that these practices must stop, cicles have fallen. Buds ovoid-cylindric to cylindric,
as the habitat itself is increasingly found to be more 1.5–2 cm long, not resinous; cataphylls with white or
important than the production of more beef and silvery fringed margins. Leaves in fascicles of 2 or 3,
wool and much of the Pinyon-Juniper woodland oc- held in a 15–20 mm long fascicle sheath, persisting 2
curs on public lands. (rarely 3) years, spreading, (15–)18–25(–30) cm long,
IUCN: LC straight and flexible, slightly twisted, 1.2–1.5 mm
wide, yellowish green or glaucous green; margins
minutely serrulate; apex short acute to acuminate;
Uses
stomata in fine lines on all surfaces. Pollen cones in
Firewood from the Pinyon-Juniper woodland is the clusters, spirally arranged, cylindrical, 3–4 cm long,
most common use both past and present. Pinyon purplish, turning dark or dull brown. Seed cones
pine wood has a higher than average heat value and solitary or in pairs, pedunculate to 3 cm or almost
sessile, soon falling after seed dispersal, spreading,
Uses
(7–)9–15(–18) cm long, narrowly ovoid or ovoid-­
oblong when closed, broadly ovoid or ovoid-cylin- Slash pine is a main source for naval stores, a term
dric with a more or less flattened base when open. used first by the English for course resin products
Seed scales thin woody, rigid, light brown proxi- used in the navy to make wooden ships waterproof
mally; apophysis lustrous brown, raised and sharply and tar the rigging. This industry is one of the oldest
keeled transversally; umbo dorsal and depressed- in the United States and has supplied huge quanti-
pyramidal, armed with a short, stout prickle. Seeds ties of resin and turpentine since colonial times. In
ellipsoid, 6–7 mm long, dark brown, with a wing 20– the past, resin was virtually the only product har-
30 mm long. Seedlings with or without a ‘grass’ stage. vested from Slash pine (the method of tapping gave
the tree its vernacular name), reaching a production 671
peak in the 1930s. After that date production sharply
Distribution
declined due to labour costs and other market forc-
SE USA, from South Carolina to Mississippi, S to es, but plantation trees are still tapped for this pur-
Florida Keys. pose up to 20 years of age, after which the timber
TDWG codes: 78 ALA FLA GEO LOU MSI SCA is harvested for the pulp industry, yielding sulfate
turpentine as a by-product. Selection of high resin
yielding trees has produced commercially available
Ecology
seedlings for this purpose and such seedlings have
Pinus elliottii is a subtropical pine growing in a been exported to many tropical countries. Detailed
warm and humid climate at low elevations. Most information on this industry is found in the book
rain falls in summer as short cloudbursts to a total Plant Resins (Langenheim, 2003). The other uses of
average of 1270 mm per year; winters are mild to the wood are mainly as roundwood for poles and
warm and dry although frosts do occur especially posts, often after treatment for conservation. Bark
on clear nights. It especially thrives in wetlands, and needles are used as a mulch in horticulture, but
where it is abundant on the sandy islands of exten- the tree itself is rarely planted in gardens.
sive swamps such as Okefenokee and the Everglades
and on pond margins and along drainages. Its roots 2 varieties are recognized:
need aerated soil, so it avoids the swamps proper. In
the Florida Keys, the variety densa occurs on karst Pinus elliottii Engelm. var. elliottii. Type: USA:
limestone, an extremely nutrient poor rock type South Carolina, J. H. Mellichamp s.n. [MO No.
derived from ancient coral reefs. Pinus elliottii can 3941460], 20 Mar 1873 (lectotype MO).
form pure stands, or mixed pine forest with P. taeda
and P. serotina. Other conifers that can occur with
Description
P. elliottii are Chamaecyparis thyoides and Taxodium
distichum; broadleaf trees are e.g. Nyssa sylvatica Leaves mostly in fascicles of 3; hypodermis with 2–3
and N. aquatica, Magnolia virginiana, and Persea cell layers; resin ducts 3–5. Seedlings developing
borbonia. The understorey is often dominated by normally, with regular internodes and branching
shrubby palms or palmettos (Sabal palmetto, Ser­ whorls.
enoa repens). The southernmost populations (var.
densa) develop a ‘grass stage’ as an adaptation to fre-
Distribution
quent ground fires. Initially, seedlings produce very
little apical growth and develop an extensive root SE USA, from South Carolina to Mississippi (coastal
system; after fire damage new buds are formed at the plain) including Florida to the Everglades.
apex of a very short stem near the ground. Photo­ TDWG codes: 78 ALA FLA GEO LOU MSI SCA
synthesis is with juvenile ‘grass’ leaves only. After
some years, a sudden apical growth elongates the
Conservation
stem rapidly without branching, lifting the growing
points above the heat of fires. At this stage normal IUCN: LC
subapical branching begins and fascicles with adult
needle leaves are formed.
 Pinus elliottii Engelm. var. densa Little & K. W. Dor-
Vernacular names
man, J. Forest. (Washington) 50: 921, f. 1, 2. 1952.
Pinus elliottii Engelm. subsp. densa (Little & Apache pine; pino real (Mexico)
K. W. Dorman) E. Murray, Kalmia 12: 23. 1982; Pinus
densa (Little & K. W. Dorman) Silba, Phytologia
Description
Mem. 7: 50. 1984. Type: USA: Florida, Henry Co.,
La Belle, 20 mi. (30 km) SE of La Belle, E. L. Little & Trees to 20–25(–27) m takk, d.b.h. to 70–90 cm.
K. W. Dorman 14033 (holotype US). Trunk monopodial, straight; bark thick, rough, scaly,
divided into long, irregular plates by wide, shal-
Pinus densa (Little & K. W. Dorman) Silba var. aus­ low fissures, dark brown, weathering grey. Branch-
672 trokeysensis Silba, Phytologia 68: 49. 1990, [“austro- es long, spreading or assurging, forming a broad,
keysensis”]. rounded, open crown. Shoots stout, very rough with
large, decurrent, persistent pulvini. Cataphylls large,
15–20 mm long, subulate, recurved, with scarious
Description
lamina and erose-ciliate margins, dark brown to
Leaves more frequently in fascicles of 2 than in 3’s; blackish. Vegetative buds large; terminal bud 20–30
hypodermis with (2–)3–4(–5) cell layers; resin ducts × 15–20 mm, ovoid-conical; lateral buds smaller, not
up to 9 per leaf. Seedlings usually developing a grass resinous. Fascicle sheaths persistent and remaining
stage, with shortened stems and crowded subtermi- long, (15–)25–35(–40) mm, reddish brown weath-
nal buds; growing stem mostly without branches ering brown to blackish. Leaves in fascicles of (2–)
initially. 3(–4), rarely 5, persisting 2–3 years, rigid or pliant,
straight or slightly drooping, (18–)20–35 cm long,
1.5–2 mm wide; margins serrulate; apex acute-pun-
Distribution
gent; leaf colour light green or glaucous. Stomata on
SE USA: in S Florida and north along the coasts to all faces of leaves, in (6–)8–15 lines on the convex
Central Florida; also on eight of the Lower Florida abaxial face and 4–8 lines on each adaxial face. Pol-
Keys. len cones ovoid-oblong to cylindrical, up to 4 cm
TDWG codes: 78 FLA long and 12 mm wide, yellowish pink, turning yel-
lowish brown. Seed cones subterminal, in pairs or
in whorls of 3–5, on thick, short, curved peduncles,
Conservation
persisting some time, leaving a few scales on branch
IUCN: LC when falling. Mature cones seemingly sessile, ovoid-
oblong, curved, with an oblique base when opened,
8–15 × 6–10 cm when open, often resinous. Seed
Pinus engelmannii Carrière, Rev. Hort., sér. 4, 3: scales oblong, straight or recurved, thick woody,
227. 1854. Type: Mexico: Chihuahua, Cosiquiriachi, with thin margins, up to 20 mm wide. Apophysis
F. A. Wislizenus 233 (lectotype MO). Fig. 224 prominently raised, transversely keeled, rhombic or
pentagonal in outline, ochraceous or light brown,
Pinus macrophylla Engelm. var. blancoi Martínez, often with dark radial lines or fissures. Umbo dor-
Anales Inst. Biol. Univ. Nac. México 15: 345. 1944; sal, large, rhombic to transverse-rhombic in outline,
Pinus engelmannii Carrière var. blancoi (Martínez) with a persistent, curved spine up to 3 mm long.
Martínez, Pinos Mexic., ed. 2: 288. 1948. Seeds obliquely ovoid, slightly flattened, 5–8 × 4–5.5
mm, light grey-brown, often with dark spots. Seed
wings ovate-oblong to obliquely ovate, 18–25 × 7–10
Etymology
mm, translucent light ochraceous.
This pine species was named after the German/
American botanist Georg(e) Engelmann (1809–
Distribution
1884), who became Director of the Missouri Botani-
cal Garden. USA: SE Arizona and extreme SW New Mexico;
Mexico: extending south from the populations in
the SW USA through the Sierra Madre Occidental
in Sonora, Chihuahua, NE Sinaloa, Durango and 68: 64. 1990; Pinus wangii Hu & W. C. Cheng subsp.
more scattered in Zacatecas, also in Nuevo León. kwangtungensis (Chun & Tsiang) Businský, Acta
TDWG codes: 76 ARI 77 NWM 79 MXE-CO MXE- Pruhoniciana 68: 11. 1999.
CU MXE-DU MXE-NL MXE-ZA MXN-SI MXN-SO Pinus kwangtungensis Chun & Tsiang var. variifolia
Nan Li & Y. C. Zhong, Novon 7 (3): 262. 1997; Pinus
wangii Hu & W. C. Cheng subsp. variifolia (Nan Li
Ecology
& Y. C. Zhong) Businský, Acta Pruhoniciana 68: 11.
Pinus engelmannii occurs on moderately dry, sum- 1999.
mer-warm open mountain slopes or plateaus at al- Pinus fenzeliana Hand.-Mazz. var. annamiensis
titudes between (1200–)1500–2700(–3000) m a.s.l., Silba, J. Int. Conifer Preserv. Soc. 7 (1): 30. 2000.
most abundantly between 2000–2500 m. It occurs Pinus orthophylla Businský, Willdenowia 34 (1): 229. 673
on poor rocky (volcanic) soils as well as on alluvial 2004.
coarse sand/gravel or loamy sand. The climate is Pinus eremitana Businský, Willdenowia 34 (1): 234.
temperate, with annual rainfall from 400–700 mm 2004.
increasing southward. Above 2000 m frost and snow
are common in winter. It is a constituent of open
Etymology
pine and pine-oak woodland, sometimes of mixed
pine forest, with e.g. P. leiophylla, P. lumholtzii and This pine species has been named after “Fenzel” the
P. pseudostrobus, on drier sites with P. cembroides collector of the type specimen.
and Juniperus sp., and usually with various species
of Quercus present.
Vernacular names
hai nan wu zhen song, hua nan wu zhen song (Chi-
Conservation
nese names for two species here united into one).
IUCN: LC
Description
Uses
Trees to 50 m tall, in many areas only to 20–30 m
Apache pine is commonly logged, but apparently tall; trunk to 1 m d.b.h. Bark on young trees and
not specifically selected as a timber tree; in most of branches smooth, thin, becoming scaly and flak-
its range it grows together with other pines. In some ing, brown, dark brown or grey-brown on trunks
areas depletion of larger trees has been observed. Its of larger trees. Branches spreading wide, forming
wood properties are similar to those of Ponderosa broad, umbrella-shaped or domed crowns. Foliage
pine and Jeffrey pine and the wood is put to simi- branches slender; young shoots pale brown, rarely
lar uses. In recent decades it has been taken more glaucous, glabrous or rarely puberulent in grooves,
often into cultivation as it apparently grows well in turning greyish brown. Buds ovoid to cylindrical,
regions with relatively mild winters and (moder- slightly resinous; cataphylls dark brown. Leaves in
ately) warm summers. Its very large, light green to fascicles of 5, held by deciduous sheaths of flimsy,
glaucous green needles are a striking feature in any brown scales; some leaves may fall independently
good size garden. before the whole bundle falls off the twig, produc-
ing fascicles with seemingly fewer than 5 leaves. Leaf
length extremely variable at least between popula-
Pinus fenzeliana Hand.-Mazz., Oesterr. Bot. Z. 80: tions, 4–18 cm long, leaves spreading or drooping,
337. 1931. Pinus parviflora Siebold & Zucc. var. slender, flexible, the shorter leaves straight or curved,
fenzeliana (Hand.-Mazz.) C. L. Wu, Acta Phytotax. 1–1.5 mm wide, glaucous green, with stomatal lines
Sin. 5 (3): 143. 1956. Type: China: Hainan Island, [?] on the two adaxial surfaces; margins minutely ser-
Fenzel 55 (holotype WU). rulate. Pollen cones in small clusters, short cylindri-
cal. Seed cones variable in size and shape, from short
Pinus kwangtungensis Chun & Tsiang, Sunyatsenia ovoid to long cylindrical, initially erect on stout
7: 113. 1948; Pinus wangii Hu & W. C. Cheng var. peduncles, becoming curved down to pendulous,
kwangtungensis (Chun & Tsiang) Silba, Phytologia (3–)5–15(–17) cm long, solitary or with 2–3 together.
 Seed scales soft woody, more or less flexible at base,
Distribution
cuneate to oblong; apophyses rhombic to oblong
(at base and apex of cone), curved or more or less S China, Guangdong (including Hainan Island),
straight, not recurved or more commonly recurved Guangxi, S. Hunan; Vietnam.
near cone base, becoming longitudinally furrowed TDWG codes: 36 CHH CHS-GD CHS-GX CHS-HN
in mature cones, ripening to yellowish brown or 41 VIE
reddish brown, weathering grey-brown; apex thin or
marginally thickened, usually upcurved; umbo ter-
Conservation
minal, small and obtuse. Seeds obovoid or ellipsoid,
10–15 mm long, wingless when dispersed, or with a IUCN: NT
674 variously rudimentary to small wing always shorter
than the seed and easily detached.
Uses
As a ‘soft wood’ pine, this species will yield excel-
Taxonomic notes
lent timber for construction, carpentry and perhaps
Using a somewhat broader species circumscription furniture making. Uses are presumably local only, as
here, I have united Pinus kwangtungensis with P. fen­ there are no great quantities of it anywhere. Resin
zeliana, a name which must then be taken up as the is tapped from trees in Vietnam and used as a glue.
earliest one validly published for the broader defined Because this species resembles P. morrisonicola from
species. The variability of this species, with character Taiwan, it has potential as a garden tree or for bonsai
states inconsistently distributed within and among culture. This species is currently not known to be in
populations, has led to the naming of many varie- cultivation.
ties and the classification of these with more than
one species. In Flora of China 4 (1999) P. fenzeliana,
P. kwangtungensis and P. wangii are kept separate, Pinus flexilis E. James, Account Exped. Pittsburgh
the two former each with two varieties. In this group 2: 27, 35. 1823. Pl. 29
of pines (section Quinquefolius, subsection Strobus)
the number of leaves per fascicle is five and remark-
Etymology
ably constant (very rarely a few fascicles with more
than five, but never with fewer). The variety varii­ The species epithet refers to the very flexible branch-
folia recently described by Nan Li and Y. C. Zhong es.
(op. cit.) seems to be based on erroneous observa-
tion: only juvenile growth should be counted as evi-
Vernacular names
dence for fewer than five needles in this subsection
with early deciduous sheaths which hold the nee- Limber pine, Rocky Mountain White pine
dles together. They normally remain with five even
after shedding the sheath scales, but exceptions are
Description
known to occur, most notably in the pinyon pines
(subsection Cembroides). Recently, Businský (2004) Trees to 10–15(–20) m tall, d.b.h. to 1–1.5 m; trunk
in a revision of Pinus subsection Strobus in Asia, straight or contorted, branching very low. Bark
described two new species, one from Hainan Island thin, rough and scaly, breaking into small, scaly
in China (P. orthophylla) and one from N Vietnam plates, brown, weathering grey. Branches ascend-
(P. eremitana) which both appear to belong here. ing or spreading, of higher orders slender, flexible,
The differences in character states tabulated in this assurgent. Shoots initially puberulent, soon gla-
paper are not distinct, but have overlapping values, brous, grey-green to grey. Cataphylls 5–10 mm long,
which may indicate varieties, but not distinct spe- subulate, scarious, brown. Vegetative buds ovoid
cies. Given the substantial range of measurements or subglobose; terminal bud 8–15 mm long; lateral
found for most of these characters within even the buds smaller, resinous. Fascicle sheaths ca. 15 mm
narrow species concepts of Businský, it is obvious long, deciduous and absent in second year’s fasci-
that these character states are variable within popu- cles. Leaves in fascicles of 5, persisting (3–)5–6 years,
lations, or even within single trees. straight or slightly curved, sometimes twisted, lax,
 675

3
4
2

Pl ate 2 9 . Pinus flexilis. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Seed cone.
 (5–)6–9 cm long, 0.8–1.2 mm wide, with remotely Pinus of western North America that can withstand
serrulate to nearly entire margins, acute to acumi- extreme conditions of climate on bare rock or scree
nate, dark green on abaxial face, glaucous to white without any other vegetation cover. On these sites it
stomatal lines on both adaxial faces. Pollen cones occurs either alone or with Pinus albicaulis and Abies
ovoid-oblong, 6–10 mm long, yellow. Seed cones lasiocarpa in the northern parts of its range and with
subterminal, solitary, in pairs or in whorls of 3(–4), Pinus aristata in the SE and P. longaeva in the SW. At
on stout, to 15 mm long peduncles falling with cone. lower altitudes it is usually only a minor component
Mature cones subpendulous, cylindrical, straight or of more diverse conifer forest in the south and with
slightly curved when closed, ovoid-cylindrical or Picea engelmannii, Pinus contorta, and Pseudotsuga
broad-cylindrical when opened, then 10–15 × 4–6 menziesii in the north. The seeds, which only have
676 cm. Seed scales parting but spreading only ca. 50° rudimentary wings, are mostly dispersed by rodents
from the axis, the proximal, infertile scales straight and birds, of the latter Clark’s Nutcracker (Nucifraga
or slightly recurved, the other scales obtrullate, columbiana, Corvidae) is the most important vector.
slightly concavo-convex, with seed cavities near
base on adaxial side; apex straight or recurved at
Uses
mid-cone; colour dark (reddish) brown. Apophysis
relatively thick woody, triangular to rhombic, with Although the wood of Limber pine is of high quality,
an obtuse apex, straight or recurved but not reflexed as a timber tree the species is of minor importance
at mid-cone, ochraceous to light brown, very resin- due to its occurrence at high altitudes, where it is
ous. Umbo terminal, transverse-triangular, obtuse, generally inaccessible and most of the trees do not
grey-brown, very resinous. Seeds broadly obovoid, grow into straight boles. Planted trees have a ten-
flattened, 10–15 × 8–10 mm, dark brown, sometimes dency to become multi-stemmed from a short base
with blackish spots. Seed wings vestigial to very and grow rather slowly. Locally, it has been used
small, reduced to a narrow strip around the distal much for construction in ‘pioneer times’, but more
part of the seed up to 5 mm wide, or absent when the recently its main uses have been restricted to car-
seed is free from the scale. pentry and furniture, as well as firewood. The spe-
cies has been introduced to Britain in 1851, but has
remained rare in gardens there and elsewhere in
Distribution
Europe. In the USA it is more common and several
W North America: Rocky Mountains from Alberta cultivars, most of which have glaucous blue needles
and British Columbia to New Mexico and Texas and and more compact habits, are being grown and are
some isolated localities to the east, mountains of SE in the horticultural trade.
California, Nevada, N Arizona, Utah; and extend-
ing into northern Mexico in scattered localities in 2 varieties are recognized:
Chihuahua, NE Sonora, Coahuila and Nuevo León.
TDWG codes: 71 ABT BRC 73 COL IDA MNT WYO Pinus flexilis E. James var. flexilis. Type: USA:
74 SDA 76 ARI CAL NEV UTA 77 NWM TEX 79 MXE- Colorado, El Paso Co., eastern flank of Pike’s Peak,
CO MXE-CU MXE-NL MXN-SO 0.25 miles north of Ruxton Park, Andresen J. W. &
[?] Barger A2125 (neotype SIU).
Ecology
Pinus flexilis E. James var. macrocarpa Engelm., in
Pinus flexilis is a conifer tree of the subalpine zone Rothrock, Rep. US Geogr. Surv., Wheeler, 6, Bot.:
in the Rocky Mountains and the ‘Basin and Range’ 258. 1879; Pinus novaemexicana P. Landry, Phyto­
region further west. In many respects its habitat re- logia 65: 479. 1989.
sembles that of Pinus albicaulis, which has a more
westerly and northerly distribution. Some isolated
Description
populations, here recognized as var. reflexa, occur
on high peaks in northern Mexico. In the Black Hills Seed scales imbricate but spreading at maturity to re-
of North Dakota P. flexilis may grow as low as 900 m lease seeds; apophyses straight or slightly incurved,
a.s.l., in the southern Rocky Mountains it can occur not recurved. Seeds without a wing when detached
at 3800 m. This species is one of several in the genus from the scale.
 Distribution Distribution
W North America: Rocky Mountains from Alberta USA: Arizona, New Mexico, SW Texas; Mexico:
to New Mexico, mountains of SE California, Ne- Chihuaha, Coahuila, Nuevo León, NE Sonora.
vada, N Arizona, Utah, isolated localities in North TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CO
Dakota, South Dakota and Nebraska. MXE-CU MXE-NL MXN-SO
TDWG codes: 71 ABT BRC 73 COL IDA MNT WYO
74 NDA NEB SDA 76 ARI CAL NEV UTA 77 NWM
Conservation
IUCN: LC
Conservation
677
IUCN: LC
Pinus gerardiana Wall. ex D. Don, in Lambert,
Pinus flexilis E. James var. reflexa Engelm., in Descr. Pinus, ed. 8°, 2: p. s.n. inter 144 et 145, t.
Rothrock, Rep. US Geogr. Surv., Wheeler, 6, Bot.: 79. 1832. Type: India: Uttaranchal, Kumaon, “Lt.”
258. 1879. Pinus reflexa (Engelm.) Engelm., Bot. Gerard s.n. [Wallich Cat. No. 6064] (lectotype K-W,
Gaz. 7: 4. 1882; Pinus flexilis E. James subsp. reflexa here designated).
(Engelm.) E. Murray, Kalmia 12: 23. 1982. Type:
USA: Arizona, Pima Co., Santa Rita Mts., E of
Etymology
Tucson, J. T. Rothrock 654 (lectotype MO).
This pine species was named after its European dis-
Pinus strobiformis Engelm. var. potosiensis Silba, coverer Lt. Gerard, who brought it to the attention of
Phytologia 68: 62. 1990. Nathaniel Wallich in Calcutta.
Pinus stylesii Frankis ex Businský, Acta Pruhon. 88:
6. 2008.
Vernacular names
Seed scales imbricate, spreading when mature; apo- Gerard’s pine, Chilgoza pine, Himalayan nut pine;
physes slightly recurved. Seeds with a wing to 5 mm chilghoza (Hindi); chiri, chujin (NW Himalaya)
or wing vestigial.
Description
Taxonomic notes
Trees to 25 m tall, but usually to 15–18 m; trunk to
In a recent publication on the genus Pinus, Businský 0.8–1 m d.b.h., usually branching low, sometimes
(2008) separated the populations of white pines in multi-stemmed. Bark smooth, hard, irregularly ex-
the northern part of the Sierra Madre Oriental, Mex- foliating with large thin plates, exposing fresh patch-
ico, as well as some outlier populations, as a distinct es, which later turn from yellow-green or pale green
species P. stylesii. This concept includes P. strobi­ to greyish white and create a multi-coloured pattern
formis var. potosiensis Silba and P. flexilis var. reflexa, on branches and a grey-white trunk; bark on lower
both from Cerro Potosí in Nuevo León. Chris Earle portion of trunk can become rough and fissured.
(www.conifers.org) has considered that it may be Main branches long, spreading and ascending,
of hybrid origin involving (originally) P. flexilis and forming a very wide, open crown. Foliage branches
P. strobiformis (= P. flexilis var. reflexa). As P. flexilis slender or stout, smooth, glabrous, yellowish green
retreated from the area in post-glacial time, while to olive-green. Buds ovoid, reddish brown, not res-
P. strobiformis seems to have increased its range, inous. Leaves in fascicles of 3, at first held by a basal
fewer and fewer ‘original’ hybrid trees remained and sheath, the scales of which are shed in the second
the genes from P. strobiformis may eventually sub- year, persistent 2–3 years, straight or curved and rig-
sume those from these earlier hybrid trees. If that id, spreading, 5–10 cm long, slender, ca. 1 mm wide,
scenario is correct, the name P. flexilis var. reflexa grey-green or dark green; stomata on the two adaxial
would be the most appropriate for this taxon. faces. Pollen cones clustered at the beginning of new
shoots, spirally arranged, 7.5–13 mm long, ovoid-­
 cylindrical. Seed cones solitary, ­lateral on shoot, in some areas to maintain stock. Animal husbandry,
sessile or short pedunculate, robust, 12–20(–23) cm grazing the pine stands, conflicts with this use as it
long, 8–11(–13) cm wide when open, oblong-ovoid, thwarts natural regeneration as well as planting ef-
with a more or less level base, falling 2–3 years after forts. The wood is used locally for light construction
maturity. Seed scales thick woody, rigid, 3.5–5 cm and carpentry. This species is comparatively rare in
long and 2–2.5 cm wide at mid-cone, spreading rela- horticultural cultivation (despite good hardiness)
tively wide, with two deep cavities on adaxial surface and mainly restricted to collections in arboreta.
that hold the seeds; apophysis strongly developed, Its bark is reminiscent of that of the Lacebark pine
reflexed, longitudinally ridged and grooved, apical- (P. bungeana), but somewhat less decorative; the
ly strongly curved near base of cone; umbo dorsal, seeds of P. gerardiana are much larger and better for
678 with a recurved or hooked, obtuse apex. Seeds large, comsumption (although the Chinese may of course
20–25 mm long, ovoid-oblong to more or less cylin- disagree on this).
dric, asymmetrical, 8–12 mm wide; wing rudimen-
tary, remaining attached to the seed scale.
Pinus glabra Walter, Fl. Carol.: 237. 1788. Type not
designated.
Distribution
E Afghanistan; China, S Xizang (Tibet); India, Jam-
Etymology
mu-Kashmir; N Pakistan.
TDWG codes: 34 AFG 36 CHT 40 PAK WHM-HP The species epithet means ‘smooth’ (glabrous, i.e.
WHM-JK without hairs) and pertains to the shoots.

Ecology Vernacular names

Pinus gerardiana grows in the mountains from about Spruce pine, Poor pine
2000 m to 3350 m above sea level. In the Himalayas
this means that this pine is restricted to valley floors
Description
between very high mountain ranges, which isolate
different populations to a certain extent. It prefers Trees to 30–35 m tall; trunk to 1 m d.b.h., straight,
dry, sunny slopes where the vegetation is more or columnar. Bark thick, compact, breaking into elon-
less open. In Afghanistan this pine is cultivated for gate, irregular, small plates and longitudinal fissures,
its edible seeds. They are evidently dispersed by exfoliating slowly, grey-brown weathering grey, on
birds, as is the case with other wingless or nearly branches in crown smooth, not flaking, grey. Branch-
wingless pine seeds, but detailed studies to identify es few, spreading to ascending, forming a rounded
the bird(s) and the role they play in the survival of to flat-topped crown in older trees. Foliage branches
this pine have not been undertaken as far as I know. slender, rough with pulvini of fallen leaf fascicles,
reddish or purplish brown, new shoots sometimes
glaucous, turning grey. Buds ovoid, 5–10 mm long,
Conservation
acute, slightly resinous; cataphylls red-brown, finely
IUCN: NT fringed. Leaves in fascicles of 2, held together in a
short sheath, straight, 4–8 cm long, rigid, slightly
twisted, 0.7–1.2 mm wide; margins minutely serru-
Uses
late; apex acute; leaf colour dark green; stomata in
The main economic use of this pine is of its edible, lines on all surfaces. Pollen cones in small clusters,
oil-rich seeds (neoza in Hindi), which are harvested spirally arranged, short cylindrical, 1–1.5 cm long,
by knocking the cones from the trees in autumn and purplish, turning brown. Seed cones solitary or in
early winter. Local mountain clans and villages own pairs, semipersistent, opening soon, spreading on
rights to the seeds and control the harvest, which is branch from short peduncles, 4–7 cm long, sym-
exported to markets on the plains of northern India. metrical or nearly so, ovoid-oblong before opening,
Trees that do not produce enough cones anymore are becoming more cylindrical when open. Seed scales
cut as firewood and new trees are planted from seeds thin woody, more or less flexible at base, without a
contrasting sealing border; apo­physis slightly raised, merits are deemed to be low and it is rarely grown
irregularly rhombic in outline or with a rounded outside arboreta and botanic gardens.
upper margin, transversely keeled; umbo central,
depressed or obtuse, unarmed or with a weak, de-
ciduous and small prickle. Seeds obovoid or near- Pinus greggii Engelm. ex Parl., in Candolle, Prodr.
ly deltoid, 5–6 mm long, seed coat rough, mottled 16 (2): 396. 1868.
brown, with a wing 12–15 mm long.
Etymology
Distribution
This species was named after J. Gregg, who collected
SE USA, Alabama, Florida, Georgia, Louisiana, Mis- the type specimen in 1848. 679
sissippi, South Carolina (coastal plain).
TDWG codes: 78 ALA FLA GEO LOU MSI SCA
Vernacular names
Gregg’s pine; pino prielo (Spanish)
Ecology
Pinus glabra is a lowland pine of the warm temper-
Description
ate climate in the SE United States, where summers
are long, hot and humid and winters mild. This Trees to 20–25 m tall, d.b.h. to 70–80 cm. Trunk
species occurs scattered in river valleys on banks monopodial, usually straight; bark thick, with deep,
of streams and in hummocks and swamps of the longitudinal fissures and rough, elongated plates,
southern Coastal Plain in acidic sandy soils, often greyish brown. Branches long, slender, spreading
with a high water table. It establishes itself in the or curved downward, not pendulous, forming a
shade of broad-leaf trees such as Magnolia, Lirio­ rounded, dense or more open crown. Shoots often
dendron, Liquidambar, Nyssa, Carya, Fagus, and multi-nodal, smooth, reddish brown to grey-brown.
Quercus and survives by overtopping them. This tol- Cataphylls lanceolate to subulate, straight, erose-
erance to shade is rather unusual for a pine and it ciliate at margins, dark brown. Vegetative buds
is diminished as it grows taller, so it often requires ovoid-conical to oval-oblong; terminal bud 10–15
some canopy opening to attain dominance. Un- mm long; lateral buds more ovoid and smaller, not
like many other pines it is susceptible to fire. It may resinous. Fascicle sheaths initially ca. 10 mm long,
occasionally grow with other pines such as Pinus persistent but reduced in mature fascicles to 3–5
echinata, P. elliottii and P. taeda or with Taxodium mm. Leaves in fascicles of 3, usually rigid, or flex-
distichum. Numerous other angiosperms are com- ible and drooping, persisting up to 4 years, straight,
ponents of these swamp forests, where waterlogging (7–)9–13(–15) cm long, 1–1.2 mm wide, serrulate
is only intermittent, most are trees and shrubs or along margins, acute-pungent, lustrous green. Sto-
woody vines. mata in several lines on all faces of the leaves. Pol-
len cones ovoid-oblong to cylindrical, 1.5–2 cm ×
5–6 mm, yellowish. Seed cones appearing on very
Conservation
young trees, subterminal or lateral, in whorls of 3–8,
IUCN: LC sometimes more, on short, stout, tenacious pedun-
cles, at maturity appearing sessile, long persistent,
serotinous. Mature cones serotinous, narrowly and
Uses
obliquely ovoid to ovoid-attenuate, slightly curved
The quality of its timber is poor, the wood is brit- or straight when closed, with an oblique base, (6–)
tle and not durable while the scattered occurence of 8–13(–15) × (4–)5–7 cm when open (width 3.5–5 cm
this species also discourages exploitation on a large when closed). Seed scales remaining closed several
scale. Locally it may support a small scale forest in- to many years, parting eventually or not, oblong,
dustry if mixed with other pines. In some parts of straight or recurved when parted. Apophyses flat
its range it is planted and harvested for Christmas or slightly raised, rhombic to obtrullate in outline,
trees, but it needs shearing repeatedly to attain the with an undulate or crenate upper margin, (weakly)
desired density of foliage and shape. Its horticultural transversely keeled, sometimes gibbous on one side
 towards base of cone, lustrous ochraceous or light seems to grow much faster in trial plots than several
brown, weathering grey. Umbo dorsal, depressed other species (Dvorak & Donahue, 1992). Gregg’s
or flat, with a minute, deciduous prickle. Seeds ob- pine is rare in cultivation and probably restricted to
liquely obovoid to ellipsoid, slightly flattened, 5–8 × botanical collections (arboreta), although in Italy it
3–4 mm, grey-brown to blackish brown. Seed wings is sometimes planted as an amenity tree.
oblong, with a straight and a curved side, 15–20 ×
5.5–8 mm, yellowish to grey-brown. 2 varieties are recognized:

Pinus greggii Engelm. ex Parl. var. greggii. Type:

Distribution
Mexico: Coahuila, San Antonio de las Alanzanas,
680 Mexico: in extreme SE Coahuila, S Nuevo León, SE near Saltillo, J. Gregg 402 (lectotype MO).
San Luís Potosí, Querétaro, Hidalgo and N Puebla.
TDWG codes: 79 MXC-PU MXE-CO MXE-HI MXE-
Description
NL MXE-QU MXE-SL
Leaves 7–12 cm long, rigid, spreading to erect, with
all resin ducts medial. Seed wings 5.5–6.4 mm wide.
Ecology
The altitudinal range of this species is 1300–2600
Distribution
m, in the northern part of its distribution 2300–
2700 m a.s.l. Annual precipitation varies between Mexico: Coahuila, Nuevo León, San Luis Potosí.
600–800 mm in much of its range, except on the TDWG codes: 79 MXE-CO MXE-NL MXE-SL
eastern escarpment of the mountain ranges along
the Hidalgo-Veracruz borderline, where it is 1000–
Ecology
1600 mm. In the north it is more often found on
slightly alkaline soils (pH 7.0–8.0), in the south on This northern variety is mainly found on alkaline
acid soils (pH 4.0–5.0) (Dvorak & Donahue, 1992). soils (pH 7–8) derived from limestone. It is most
It is nowhere abundant in its scattered range, and commonly found scattered in mixed pine-oak ­forest.
always occurs mixed with e.g. Quercus, Platanus,
Liquidambar, and Fraxinus, other pines, e.g. Pinus
Conservation
patula, P. pseudo­strobus, P. teocote, P. montezumae,
and P. arizonica var. stormiae, with P. cembroides and According to extent of occurrence (EOO) data this
Juniperus flaccida on dry sites, and at higher and variety rates as Vulnerable, but according to its area
more mesic locations with Abies vejarii, Pseudotsuga of occupancy (AOO) it is possibly Near Threatened.
menziesii, or Cupressus lusitanica. The serotinous Indications of exploitation and other human impact
cones indicate adaptation to fire, but no studies on justify it to be NT. These impacts appear to be less
how this affects seed dispersal and germination have severe than in the area of the southern variety. No
been undertaken (or published). collections were recorded from protected areas.
IUCN: NT
Uses
Pinus greggii Engelm. ex Parl. var. australis
Although locally exploited with other pines, Gregg’s Donahue & Lopez, Sida 18 (4): 1092. 1999. Type:
pine is not specifically in demand as a timber tree Mexico: Queretaro, Landa de Matamoros, El
in Mexico. In many areas it has been severely de- Madroño, J. K. Donohue & J. Lopez Upton B30
pleted by general logging and overexploitation of (holotype MO).
forests. Foresters from abroad are taking an inter-
est in its potential as a forest plantation tree in other
Description
countries; it has been introduced for that purpose
in (among other countries) India, South Africa, Leaves 10–15 cm long, flexible, frequently drooping,
Zimbabwe, Argentina, and Brazil. Like another, and sometimes with 1–2 internal resin ducts. Seed wings
probably related, “closed-cone” pine, P. radiata, it 6–8 mm wide.
 seed scales have rounded apophyses terminating in
Distribution
a spiny umbo and the seeds have short wings.
Mexico: Querétaro, Hidalgo, Puebla.
TDWG codes: 79 MXC-PU MXE-HI MXE-QU
Distribution
Japan: Central and N Honshu, S Hokkaido.
Ecology
TDWG codes: 38 JAP-HN JAP-HK
This variety is generally growing on acid soils. It is
a relatively rare pine that always grows sporadically
Ecology
among other species of pine, or in mixed pine-oak
woodland. This nothospecies occurs in subalpine regions of 681
S Hokkaido and Central and N Honshu in scrubby
vegetation at altitudes between 1000 m and 1700 m
Conservation
a.s.l.
The range of this variety, based on herbarium collec-
tions data, is just outside the threshold for VU (EOO
Conservation
= 21,155 km2). Change in land use especially conver-
sion to agriculture is widespread in the region and IUCN: LC
almost certainly affects both EOO and AOO of this
taxon. The subpopulations of this variety are often
some distance apart and surrounded by agricultural Pinus halepensis Mill., Gard. Dict., ed. 8: Pinus No.
land. Grazing may have a significant effect on regen- 8. 1768. Type not designated.
eration, because areas that are not grazed showed
regeneration as opposed to those where grazing was Pinus ceciliae Llorens & L. Llorens, Folia Bot. Misc. 4:
evident. Some locations are within protected areas: 55. 1984; Pinus halepensis Mill. var. ceciliae (Llorens
the Sierra Gorda, Los Marmoles and Cuenca Hidro- & L. Llorens) Roselló et al., Candollea 47 (1): 67. 1992.
grafica del Rio Necaxa Reserves.
IUCN: VU (A2, c, d)
Etymology
The species epithet is derived from Aleppo [Haleb]
Pinus ×hakkodensis Makino, in Makino & Nemoto, on the coast of Syria.
Fl. Japon., ed. 2: 148. 1931. Pinus pentaphylla Mayr
var. hakkodensis (Makino) Kusaka, in Iwata &
Vernacular names
Kusaka, Conif. Jap. Ill. ed. 2: 151. 1954. Type not
designated. Aleppo pine; pino de Alepo, pino carrasco (Span-
ish); pin d’Alep (French); pino d’Aleppo (Italian)
Etymology
Description
The epithet means ‘from Hakkoda’, which is an or-
thographic variant of Hokkaido. Trees to 20 m tall; trunk to 80 cm d.b.h., straight,
curved or sinuous, often forked. Bark thin, grey-
brown, becoming thick only on lower trunk of large
Vernacular names
trees, then fissured longitudinally, scaly, breaking
Hakkoda-goyo (Japanese) into elongated plates, purplish brown to red-brown,
weathering grey. Branches numerous, spreading and
A natural hybrid between Pinus parviflora var. pen­ ascending, forming a broadly rounded, sometimes
taphylla and P. pumila, with characters more or less flat-topped, dense crown. Foliage branches slender,
intermediate between those of the parent species. It 2–3 mm thick (to 6 mm on cone-bearing shoots),
is a shrub with ascendent stems to 3 m tall. The seed nearly smooth, glabrous, new shoots glaucous or
cones are ellipsoid-ovoid, ca. 6 cm long and 3.5 cm grey-green, becoming ash-grey. Buds ovoid-­conical,
wide and open at maturity to release the seeds. The acute, 5–10 mm long, not resinous; cataphylls with
 recurved apices, reddish brown fringed with white TDWG codes: 12 BAL COR FRA-FR FRA-MO SAR
hairs. Leaves in fascicles of 2, held by a 9–12 mm SPA-SP 13 ALB GRC ITA-IT SIC-MA SIC-CI YUG-CR
long, persistent sheath, falling in the third year, 20 ALG LBY MOR-MO MOR-SP TUN 34 LBS-LB LBS-
straight and rigid, spreading, (6–)7–10(–12) cm SY PAL-IS
long, 0.7–0.8 mm wide; margins minutely serru-
late; leaf colour light green; stomata in fine lines on
Ecology
all surfaces. Pollen cones in clusters at base of new
shoots, spirally arranged, short cylindrical, 1–2 cm Aleppo pine grows in the hotter parts of the Medi-
long, yellow. Seed cones solitary or in whorls of 2–3, terranean coast, where brush and forest fires are
short pedunculate, directed backward or more or frequent. Despite this, its seed cones are only semi-
682 less spreading when full grown, long persistent, nar- serotinous and do open in the absence of fire in the
rowly or broadly ovoid-conical when closed, some- heat of the sun. Although closed stands exist, it is
times slightly asymmetrical, 6–10(–12) cm long, more commonly scattered in maquis or garrigue
variously serotinous, 3–5 cm wide when closed, 4–7 vegetation on sunny hills and slopes down to the
cm wide when opened. Seed scales thick woody, rig- sea shore, most commonly on limestone and dolo-
id, straight, oblong; apophyses nearly flat or slightly mite. In stands where fire has been absent for a
raised, weakly transversely keeled and with thin longer period, oaks (Quercus suber, Q. ilex) invade
rays radiating from the centre, more or less rhombic and will eventually dominate. Presumably increased
or often with a rounded upper margin, to 20 mm frequency of fire caused by human activities gives
wide at mid-cone, lustrous orange-brown or red- the advantage to P. halepensis. Its altitudinal range is
brown weathering grey; umbo flat or depressed, 4–7 from sea level to ca. 1700 m (in Morocco).
mm wide, broadly rhombic in outline, tan or grey-
brown, unarmed. Seeds obovoid, slightly flattened,
Conservation
(5–)6–7(–8) mm long, grey-brown, dark mottled;
wing 15–25 mm long, 8–12 mm wide, more or less IUCN: LC
straight, grey-brown with darker streaks.
Uses
Taxonomic notes
The wood of Aleppo pine, due to size and shape
Varieties described of Pinus halepensis by some au- of most trees and poor quality, is of little value as
thors are here considered to be varieties of P. brutia. timber. It is presently used as firewood and for
Pinus brutia and P. halepensis are closely related and charcoal burning, in the past it served for mine
the first is sometimes treated as a variety or subspe- props, raiway sleepers, and telephone poles. It is
cies of the latter. Pinus halepensis has somewhat rich in resin and still tapped locally for that prod-
shorter and certainly thinner needles, not reach- uct; plantations have been established for this use in
ing 1 mm diam.; its cones are more elongate when Greece. Witches brooms are common in this pine
closed and tend to be deflected back on the branch and could be a source of dwarf cultivars in Mediter-
by a curved peduncle, instead of spreading. Unlike ranean countries; the species is generally not hardy
P. brutia, P. halepensis is remarkably constant in in more northern latitudes. It has been planted as
its morphological characters throughout its wide wind breaks and to stop soil erosion on dry slopes.
range; this could be an indication, to be tested with Ill considered plantations established in the south-
research into its DNA, that it originated from a par- ern hemisphere have given rise to invasiveness of
ticular population of P. brutia in the eastern Medi- this species; in Australia, New Zealand, and South
terranean and quickly spread westward upon the Africa P. halepensis is now a serious weed threaten-
advent of substantial climate change. ing natural and often species-rich vegetation as well
as land used for agriculture.
Distribution
Mediterranean, from Morocco and Spain to Greece
and the coast of Libya at Jabal al Akhdar, and in Is-
rael, Lebanon, SW Syria.
Pinus hartwegii Lindl., Edwards’s Bot. Reg. 25: (weakly) transversely keeled, sometimes gibbous,
62. Aug 1839. [Allg. Gartenzeitung 7: 324. 1839]. rhombic in outline with an angular or irregular up-
Type: Mexico: locality unknown, C. T. Hartweg s.n. per margin, varying from light brown to purplish
(lectotype P). Fig. 225 brown with a blackish centre. Umbo dorsal, de-
pressed, flat or raised, obtuse or with a deciduous
Pinus rudis Endl., Syn. Conif.: 151. 1847; Pinus mon­ prickle. Seeds broadly ovoid, slightly flattened, 5–6
tezumae Lamb. var. rudis (Endl.) Shaw, [Pines Mex- mm long, light or dark brown with black spots. Seed
ico] Publ. Arnold Arbor. 1: 22. 1909; Pinus hartwegii wings oblong with a straight and a curved side, 12–
Lindl. var. rudis (Endl.) Silba, Phytologia 68: 50. 20 × 7–12 mm, brown tinged with dark stripes.
1990.
Pinus donnell-smithii Mast., Bot. Gaz. 16: 199. 1891. 683
Distribution
Mexico: in Chihuahua (Cerro Mohinora), S Coahu-
Etymology
ila, S Nuevo León, Durango, SW Tamaulipas, Jalisco
This species was named after the plant collector (Nevado de Colima), Michoacán, México, More-
C. T. Hartweg, who made extensive collections in los, Hidalgo, Distrito Federal, Tlaxcala, Puebla,
Mexico in the decade 1838–1848. W Veracruz, Guerrero (Cerro Teotepec and vicin-
ity), Oaxaca and Chiapas; in Guatemala in most of
the SW highlands; in Honduras on several isolated
Vernacular names
mountain summits; reported from the extreme N of
Hartweg’s pine; ocote, pino de las alturas (Mexico) El Salvador, but not confirmed.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
PU MXC-TL MXE-AG MXE-HI MXE-QU MXE-SL
Description
MXE-ZA MXG-VC MXN-SI MXS-GR MXS-JA MXS-MI
Trees to 25–30 m tall, d.b.h. to 0.0–1 m; trunk mo- MXS-NA MXS-OA MXT-CI 80 GUA HON
nopodial, erect. Bark thick, scaly, deeply fissured, di-
vided into small or large, square or irregular plates;
Ecology
outer bark dark brown, weathering grey. Branches
spreading and assurging, often persistent, forming a Pinus hartwegii is the typical high altitude pine of
dense, rounded crown in mature trees. Shoots thick, Mexico and Guatemala, where it often forms ex-
rigid, purplish brown, sometimes glaucous. Cata- tensive, monotypic pine forests up to the tree line
phylls 15–20 mm long, subulate, recurved, scarious, on high, isolated volcanos or summits of mountain
brown, weathering blackish grey. Vegetative buds ranges. In Honduras it is rare, of limited extent and
ovoid-conical; terminal bud up to 30 × 15 mm; lat- usually found with Abies guatemalensis, Cupres­
eral buds smaller, not resinous. Fascicle sheaths of sus lusitanica, Juniperus standleyi, Quercus spp.,
young leaves 30–40 mm long, reduced in older fasci- Dendropanax lempirianus, Drymis granadensis,
cles to (10–)15–20 mm, resinous. Leaves in fascicles a ground cover of Ericaceae, Lycopodiaceae, and epi-
of (3–)4–5(–6), most commonly 5, persisting 2(–3) phytic Bromeliaceae in a cool cloud forest type usu-
years, straight or curved, not twisted, rigid, (6–)10– ally between 2700–2850 m on the highest mountain
17(–22) cm long, (1–)1.2–1.5 mm wide, with serrulate summits. Similar forests occur in Guatemala and the
margins, acute-pungent, dull light green or glaucous southern states of Mexico, but there extensive pine
green. Stomata on all faces of leaves. Pollen ovoid- forests predominate, in which P. hartwegii increas-
oblong to cylindrical, to 20–25 × 6–7 mm, yellow- ingly dominates with rising altitude. Its altitudinal
ish or pink-purplish, maturing to brown. Seed cones range in Guatemala and Mexico is similar: (2300–)
subterminal, in pairs or whorls of 3–6 on short, stout 2500–4000(–4300) m a.s.l. At lower elevations it is
peduncles which are hidden by basal scales. Mature often mixed with P. montezumae, with which it is
cones ovoid-oblong, with acute, curved apex when closely related, and with other pines depending on
closed, (obliquely) ovoid with a flattened base when the geographical area. Soils are both from volcanic
opened, 8–12(–14) × 5–8 cm. Seed scales spreading and granitic rock, of various depths, but often poor
wide, often recurved near base, oblong, thin woody, in nutrients. Climatically there are considerable dif-
flexible or more rigid. Apophysis more or less flat, ferences congruent with latitude/altitude gradients,
 with heavy frost and snow during several months and ascending, forming a conical crown, in old trees
and often high winds near the tree lines of the high rounded or irregular. Foliage branches stout, slightly
volcanos in Central Mexico. rough after leaves have fallen, new shoots glabrous,
light brown and glaucous or pruinose, turning brown
and finally grey. Buds ovoid-oblong, 10–15 mm long,
Conservation
acute, not resinous; cataphylls papery, fringed white
IUCN: LC towards apex. Leaves in fascicles of 2, held in a short,
persistent basal sheath, spreading in stiff pseudo-
whorls, remaining on branches 5–6 years, 6–11 cm
Uses
long, rigid, straight or more often curved, 1.2–1.5
684 Pinus hartwegii has fairly dense, but resinous wood. mm wide, lustrous green or dark green; margins
It is exploited as a timber tree where stands are ex- minutely serrulate; apex acute; stomata in fine lines
tensive and accessible, but due to high altitude and on all surfaces. Pollen cones in large clusters near
consequent lack of infrastructure (access roads) base of new shoots, spirally arranged, ovoid-­oblong
many forests remain virtually untouched today. Its becoming cylindrical at anthesis, 2–3 cm long,
wood is used as round wood for posts and sawn bright yellow. Seed cones solitary or in whorls of 2–3
for construction or railway sleepers, while it is also on short peduncles, ovoid or narrowly ovoid when
pulped for the paper industry. Despite its hardiness, closed, dark green or bluish purple, soon opening
originating from very high altitudes, it is very rare in and turning dull light brown to grey, 6–10 cm long,
cultivation, in some instances under its taxonomic 4–6 cm wide when open, with a more or less flat and
synonym P. rudis. asymmetrical base. Seed scales thin woody, soft and
flexible at their base, oblong, spreading wide, light
brown; apophyses nearly flat or (slightly) raised,
Pinus heldreichii H. Christ, Verh. Naturf. Ges. rhombic to broadly rhombic, transversely keeled;
Basel 3: 549. 1863. Type not designated. Fig. 226 umbo depressed or raised, armed with a minute,
deciduous prickle. Seeds elliptic, (4–)5–6(–7) mm
Pinus leucodermis Antoine, Oesterr. Bot. Z. 14: 366. long, brown; wing 15–25 mm long, 5–7 mm wide,
1864; Pinus heldreichii H. Christ var. leucodermis oblique at the distal end.
(Antoine) Markgr. ex Fitschen, in Beissner, Handb.
Nadelholzk., ed. 3: 404. 1930; Pinus heldreichii
Taxonomic notes
H. Christ subsp. leucodermis (Antoine) E. Murray,
Kalmia 13: 23. 1983. Pinus leucodermis Ant. is still often treated as ­either
a distinct species, or a variety of P. heldreichii, but
the stated differences between the two, such as col-
Etymology
our of shoots, buds, bark, needles, and cones are not
This pine species was named after the German bota- only trivial and rather variable, they are inconsist-
nist T. H. H. von Heldreich (1822–1902). ent, with herbarium specimens and/or trees some-
times displaying character states that ‘belong’ to
the other species. The seed cones, apart from the
Vernacular names
alledged differences in colour (green versus purple
Heldreich’s pine, Bosnian pine in the growing stage, a trait found in many popula-
tions of several species in Pinaceae and usually not
of taxonomic significance) are so similar and share
Description
uniquely soft scales not found in other species in
Trees to 25(–30) m tall; trunk to 2 m d.b.h., some the section Pinus, that they must belong to a single
old trees with squat boles. Bark thick, dense and species. They are here treated as synonyms, there-
hard, breaking into numerous ridges and small an- fore the earlier name P. heldreichii H. Christ applies.
gular plates separated by shallow, irregular fissures, Molecular data indicate that P. heldreichii is not
grey-brown weathering grey. Branches spreading closely related to P. nigra, but belongs to a group of
Mediterranean pines instead. Putative hybrids with Pinus henryi Mast., J. Linn. Soc., Bot. 26: 550.
P. ­nigra found in the Balkans (see Vidaković, 1991, 1902. Pinus massoniana Lamb. var. henryi (Mast.)
citing work by P. Fukarek in Serbo-Croatian) may C. L. Wu, Acta Phytotax. Sin. 5 (3): 153. 1956; Pinus
therefore be somewhat doubtful. These authors also tabuliformis Carrière var. henryi (Mast.) C. T. Kuan,
­reported successful controlled hybridization be- Fl. Sichuan. 2: 113. 1983; Pinus tabuliformis Carrière
tween Bosnian pine (P. heldreichii var. leucodermis) subsp. henryi (Mast.) Businský, Acta Pruhoniciana
and European black pine (P. nigra subsp. nigra). 68: 26. 1999. Type: China: Hubei, [locality un-
known], A. Henry 6909 (holotype K).
Distribution
Pinus massoniana Lamb. var. wulingensis C. J. Qi &
Balkan Peninsula, scattered from Bosnia Herzego­ Q. Z. Lin, Bull. Bot. Res. Harbin 8 (3): 143. 1988. 685
vina to N Greece, Bulgaria (Rhodope Mts.) and S
Italy (Calabria and Brasilicata).
Etymology
TDWG codes: 13 ALB BUL GRC ITA-IT YUG-BH
YUG-CR YUG-MA YUG-SE This pine species was named after Augustine Henry
(1857–1930), an English dendrologist who was an
early botanical visitor to China.
Ecology
Pinus heldreichii is a montane to subalpine species,
Vernacular names
in Albania, Bosnia and Serbia usually growing above
2200 m a.s.l. up to 2640 m, but elsewhere down to Henry’s pine
1000 m. It is most commonly found on steep moun-
tainsides with very thin soil over limestone, most of-
Description
ten in pure, scattered stands. It grows very slowly in
this habitat and presumably ancient specimen trees Trees to 25 m tall; trunk to 1 m d.b.h., monopodial.
are known, e.g. in Calabria. Although occurring in Bark scaly, fissured, breaking into large irregular
the Mediterranean region, its altitudinal range sub- plates, greyish brown weathering grey, flaking in
jects it to sometimes quite severe winter frosts. small or large chips. Branches spreading or curving
down, contorted, forming a domed or flat-topped
crown. Foliage branches stout, becoming rough with
Conservation
pulvini after shedding leaves, glabrous, new shoots
IUCN: LC reddish brown, usually glaucous, turning light
brown to grey-brown or grey. Buds oblong, acute;
terminal bud to 20 mm long, slightly resinous; cata-
Uses
phylls appressed, pale brown. Leaves in fascicles of
Heldreich’s pine is not an important timber tree, al- 2, more or less remote, persisting 2–3 years, spread-
though it is planted in some Balkan countries with ing, held in a 10–15 mm long, persistent basal sheath,
timber production as an objective. The wood is used 7–12 cm long, straight or curved, pliant, 0.7–1 mm
for heavy construction or as round timber, e.g. for wide, often slightly twisted, green; margins minutely
poles, and to build traditional fences. Its main value serrulate; apex acute or acuminate; stomata in fine
is as an ornamental tree and it is fairly widely cul- lines on all surfaces. Pollen cones in clusters at base
tivated for this purpose. It forms as a young tree a of new shoots, spirally arranged, short cylindrical,
dense, conical crown and has attractive glaucous to ca. 2 cm long, yellow. Seed cones solitary or in pairs,
light grey shoots and dark green needles. In culti- short pedunculate, usually persisting a few years but
vation it grows relatively fast, given better condi- opening soon, ovoid when closed, near-symmetri-
tions than in its natural habitat. Several cultivars are cal, 2.5–5 cm long and 2.5–5 cm wide when opened.
known, usually under the synonym P. leucodermis. Seed scales thin woody, rigid, oblong to obovate,
Some of these are slow growing ‘dwarf ’ conifers. except basal scales spreading wide, often recurving,
In the Balkans, a number of botanical varieties has brown. Apophyses slighty raised, variously shaped,
been named under P. heldreichii, it is not known if transversely keeled or with 4 radial ridges, ripen-
any of these are in cultivation. ing from green to lustrous brown or reddish brown;
 umbo depressed with a minute prickle. Seeds ovoid- Pinus herrerae Martínez, Anales Inst. Biol. Univ.
oblong, 3.5–5 × 2–3 mm, pale brown, often mottled; Nac. México 11: 76. 1940 [“herrerai”]. Pinus
wing 9–12 mm long, 4–5 mm wide, dark brown. teocote Schiede ex Schltdl. & Cham. var. herrerae
(Martínez) Silba, Phytologia 68: 63. 1990. Type:
Mexico: Jalisco, Tecalitlán, Sierra del Halo,
Taxonomic notes
M. Martínez 3427 (lectotype MEXU).
This species has been treated as a variety of Pinus
tabuliformis in Flora of China 4 (1999) and is indeed
Etymology
similar in many characters to that species. Its leaves
are not wider than 1 mm and its seed cones are small- This species was named after “Prof. D. Alfonso
686 er, with scales that have only slightly raised apophys- L. Herrera”, a Mexican naturalist.
es. It was also classified as a variety of P. massoniana,
from which it differs in its much shorter leaves, its
Vernacular names
uninodal growth of shoots, smaller, nearly globose-
ovoid seed cones, and relatively shorter seed wings. Herrera’s pine; Ocote, Pino chino (Spanish)

Distribution Description
China: Chongqing, Hubei, Hunan, Shaanxi, Si- Trees to 30–35 m tall, d.b.h. to 0.8–1 m; trunk mo-
chuan. nopodial, straight or sometimes tortuous. Bark
TDWG codes: 36 CHC-CQ CHC-HU CHC-SC CHN- thick, rough, with scaly plates and shallow, longitudi-
SA CHS-HN nal fissures, reddish brown to grey-brown. Branches
long, slender, arching or spreading horizontally, the
foliage slightly pendulous. Shoots smooth, with well
Ecology
developed, long decurrent pulvini, orange-brown.
Pinus henryi is most common at middle elevations Cataphylls subulate, scarious, recurving, brown.
in the mountains of Central China, and occurs from Vegetative buds ovoid-acute to ovoid-oblong or cy-
1100 m to 2000 m a.s.l. It prefers dry, sunny slopes lindrical; terminal bud 10–15 mm long; lateral buds
and hills, where competition from broad-leaved smaller, not resinous. Fascicle sheaths up to 20 mm
trees is less severe as the woods are more open and long, torn by elongating and spreading leaves, per-
lower than in more mesic sites. It is also a pioneer in sistent but reduced to 8–15 mm in mature fascicles.
secondary vegetation, and is there commonly mixed Leaves in fascicles of 3, persisting 3 years, slender,
with deciduous shrubs and trees, in later stages of the lax, drooping or spreading, (10–)15–20 cm long,
succession often giving way to these angiosperms. 0.7–0.9 mm wide, serrulate at margins, acute, light
green to yellowish green. Stomata on all faces of
the leaves. Pollen cones spreading, ovoid-oblong
Conservation
to cylindrical, 1.5–1.8 cm × 5 mm, yellowish green,
IUCN: NT tinged red. Seed cones subterminal, solitary or op-
posite, rarely in whorls of 3, on 10–15 mm long pe-
duncles, at first erect, soon on recurved peduncles
Uses
which fall with cones. Mature cones narrowly ovoid
This species is similar in its wood properties to when closed, nearly symmetrical or slightly curved,
P. tabuliformis, but it is less common and wide- ovoid when opened, (2–)3–3.5(–4) × 2–3.5 cm when
spread, and as a consequence of this less important open. Seed scales thick woody, oblong, straight or
as a timber tree. Pinus henryi is apperently absent recurved. Apophysis slightly raised, mostly so on a
or very rare in cultivation; it was not introduced to few proximal scales on one side of cone, transversely
Europe when the opportunity arose during the early keeled; apical margin entire or crenate, radially stri-
decades of the 20th century. ate, light brown. Umbo dorsal, pyramidal, with a
minute, ­deciduous prickle. Seeds (ob)ovoid, slightly Pinus hwangshanensis W. Y. Hsia, Contr. Inst. Bot.
flattened, 2.5–4 × 2–3 mm, dark grey-brown. Seed Natl. Acad. Peiping 4: 155. 1936. Pinus luchuensis
wings obliquely ovate, 5–8 × 3–5 mm, translucent, Mayr var. hwangshanensis (W. Y. Hsia) C. L. Wu,
yellowish with a dark tinge. Acta Phytotax. Sin. 5 (3): 158. 1956; Pinus luchuensis
Mayr subsp. hwangshanensis (W. Y. Hsia) D. Z. Li,
Edinburgh J. Bot. 54 (3): 341. 1997. Type: China:
Distribution
Anhui, Huang Shan, Si Hai Men, P. C. Tsoong 3111
Mexico: along the Sierra Madre Occidental and (holotype PE).
more abundantly along the Sierra Madre del Sur; in
SW Chihuahua, Sinaloa, Durango, W and S Jalisco, Pinus taiwanensis Hayata var. damingshanensis
Michoacán and Guerrero. W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): 687
TDWG codes: 79 MXE-CU MXE-DU MXN-SI MXS- 85. 1975.
GR MXS-JA MXS-MI
Etymology
Ecology
The species epithet hwangshanensis refers to the
The altitudinal range of P. herrerae is (1100–)1500– Huang Shan mountain range in S Anhui, China.
2600 m a.s.l.; its lowest altitude apparently is reached
in the Sierra de Cuale in W Jalisco. It occurs in the
Vernacular names
mesic forest belt, with annual precipitation of ca.
900–1600 mm, but with a dry season lasting from Huang Shan pine; huang shan song (Chinese)
November to May. From N to S annual precipitation
increases. Mixed pine and pine-oak forest is its usu-
Description
al habitat, in which it is associated with many other
pines according to their ranges, further with Pseudo­ Trees to 45 m tall; trunk to 1 m d.b.h, bole straight or
tsuga locally. Other broad-leaved trees, e.g. Arbutus, more or less tortuous. Bark on trunk rough and scaly,
Alnus, Clethra, Juglans, Persea, Clusia, and Tilia, are breaking into large plates and deep fissures, grey-
sometimes common. In many areas the forests are brown to dark grey. First order branches long and
greatly influenced by man-made changes in their spreading, slowly and sometimes only partly self-
composition, e.g. by selective felling of larger trees pruning, leaving stumps on the trunk; higher order
of Pinus, or by recurrent fires and grazing. branches assurgent and densely crowded, forming
flat-topped or domed crowns in natural habitat. Fo-
liage branches glabrous, more or less smooth, light
Conservation
yellowish brown. Buds ovoid-conical to cylindri-
IUCN: LC cal, 10–15 mm long, 5–7 mm wide, resinous; cata-
phylls appressed, reddish or chestnut brown. Leaves
in fascicles of 2, held by a persistent, slender basal
Uses
sheath 5–10 mm long, straight or slightly curved,
Along with other tall growing pines, this species is (5–)10–17 cm long, slender, pliant, slightly twisted,
(heavily) exploited for timber throughout almost 0.6–1 mm wide; margins minutely serrulate; apex
all of its range. It is considered to be of good wood acute; stomata in fine lines on all surfaces. Pollen
quality. Provenance sampling has been started and cones clustered, spirally arranged, short cylindri-
is recommended by foresters (Dvorak & Donahue, cal, 1.5–2 cm long, yellow tinged with red, becoming
1992) to be conducted on a larger scale. This pine is reddish brown. Seed cones solitary or sometimes in
unknown in horticulture. pairs, persistent on short peduncles, spreading, 3–6
cm long, narrowly ovoid when closed, widening to
2.5–5 cm when open. Seed scales thin woody, rigid,
oblong, ca, 2.5 × 1.3 cm at mid cone (in larger cones),
straight when spreading, chocolate brown. Apophy-
sis rhombic in outline or with rounded upper mar-
gin, nearly flat, transversely keeled, slightly ­rugose,
 lustrous light brown; umbo broadly ellipsoid, de- outcrops, but in many areas forest disturbance has
pressed, armed with a small, persistent prickle. probably favoured the light demanding pines over
Seeds ellipsoid-ovoid, 5–6 mm long, slightly flat- broad-leaved trees.
tened; wing 15–20 mm long, persistent.
Conservation
Taxonomic notes
IUCN: LC
The variety Pinus taiwanensis var. damingshanensis
W. C. Cheng & L. K. Fu was described from main-
Uses
land China (Guangxi, Daming Shan), with both
688 marginal and medial resin ducts. Needles of the This pine has good quality timber with suitable
same species in Taiwan are reported to have medial, strength for construction, e.g. in buildings and
but rarely also marginal resin ducts. Probably this wooden bridges, use as railway sleepers and mine
population, growing on a single mountain, belongs props. Its wood is also used for building fences and
to P. hwangshanensis W. Y. Hsia. This species is very gates, crates and boxes, panelling, flooring, furniture
similar to both P. luchuensis and P. taiwanensis, but making, industrial and domestic woodware, tools,
differs from them in having a persistent, mucronate plywood, fibreboard, and wood pulp. Forestry plan-
prickle on a depressed seed scale umbo. The position tation of this species (and its varieties) is restricted
of resin ducts is a variable character in this group of to SE China as far as is known. In horticulture it is
species and less reliable. The buds of P. hwangshan­ rare, except for the use for bonsai growing popular
ensis are darker brown than those of P. taiwanensis. in E Asia. It may occur in arboreta under the name
Other characters, e.g. colour of pollen cones, need P. taiwanensis, but if these trees originated from
further verification based on multiple observations mainland China they are more likely P. hwangshan­
to see whether they hold true. I concur with Mill’s ensis.
observation in Flora of China 4: 17 (1999) and here
restrict P. taiwanensis to the island of Taiwan.
Pinus jaliscana Pérez de la Rosa, Phytologia 54:
290. 1983. Pinus patula Schiede ex Schltdl. & Cham.
Distribution
var. jaliscana (Pérez de la Rosa) Silba, Phytologia
China: Anhui, Fujian, Guangxi, Guizhou, S Henan, 58: 368. 1985. Type: Mexico: Jalisco, Sierra del
Hubei, Hunan, Jiangsu, Jiangxi, SE Yunnan, Zhe- Cuale, near road ca. 4 km N of El Tuito to Zimapan
jiang. mine, 7 km from Hwy. 200, J. A. Pérez de la Rosa
TDWG codes: 36 CHC-GZ CHC-HU CHC-YN CHS- 370 (holotype IBUG).
AH CHS-FJ CHS-GX CHS-HE CHS-HN CHS-JS CHS-
JX CHS-ZJ Pinus macvaughii Carvajal, Phytologia 59: 139. 1986;
Pinus oocarpa Schiede ex Schltdl. var. macvaughii
(Carvajal) Silba, Phytologia 68: 57. 1990.
Ecology
Pinus hwangshanensis occurs in mixed montane de-
Etymology
ciduous forests, in open areas on slopes and ridges.
This forest type is known in China as Mixed Meso- The species epithet refers to the Mexican State of
phytic Forest (Wang, 1961) and occurs generally to Jalisco, where it was found.
the north of evergreen broad-leaved forests, with a
wide transitional zone. It is also prevalent at higher
Vernacular names
altitudes further south, where the evergreen forest
in the valleys and plains has all but disappeared due Jalisco pine
to agriculture. Its altitudinal range is approximately
between 500 m and 2500 m a.s.l. and the most com-
Description
mon angiosperm (broad-leaved) trees belong to the
Fagaceae. In undisturbed forest the pines would oc- Trees to 25–30(–35) m tall, d.b.h. to 60–80(–100)
cupy areas with shallow, sandy acidic soils and rocky cm; trunk monopodial, straight. Bark moderately
thick, rough, scaly with irregular, longitudinal plates tion. Pinus jaliscana is a local constituent of pine
and shallow fissures, reddish to greyish brown. or pine-oak forests with a limited extent in most
Branches spreading or ascending; branches of high- localities known so far. More common and wide-
er orders slender, flexible, slightly pendulous, form- spread species that grow with it are P. maximinoi
ing a rounded, rather open crown. Shoots smooth, and P. oocarpa, and at slightly higher altitudes also
orange-brown, later grey-brown. Cataphylls small, P. douglasiana. Several species of Quercus are often
subulate, recurving at apex. Vegetative buds ovoid- codominant, Clusia salvinii is also common.
oblong to conical; terminal bud 10–15 mm long; lat-
eral buds ovoid-acute, smaller, not resinous. Fascicle
Conservation
sheaths up to 15 mm long, light brown, persistent but
reduced to 8–10 mm on mature fascicles, weathering IUCN: NT 689
light grey. Leaves in fascicles of (4–)5, rarely 3, per-
sisting 2–3 years, slender, lax, erect or slightly droop-
Uses
ing, 12–18(–22) cm long, (0.5–)0.6–0.8 mm wide,
with serrulate margins, acute, light green to yellow- No particular preference to single this species out
ish green. Stomata on all faces of leaves, sometimes as a source of timber is known to occur; it is locally
inconspicuous. Pollen cones ovoid-oblong to cylin- exploited with other pines that grow with it and are
drical, 1.2–1.8 cm × 5–6 mm, purplish yellow, turn- more abundant. This species is not known to be in
ing light brown. Seed cones subterminal, solitary or cultivation.
in whorls of 2–3 on stout, curved, 7–15 mm long pe-
duncles remaining attached to fallen cones. Mature
cones ovoid-oblong to ovoid-attenuate when closed, Pinus jeffreyi Balf., in Murray, Bot. Exped. Oregon
often oblique at base, (4.5–)6–8.5(–9.8) cm long, (3–) 8: 2. 1853. Type: USA: California, Shasta Co., Shasta
4–5(–6) cm wide when open. Seed scales oblong, Valley, (“Chastey Valley, Lat. 41° 30´ N”), J. Jeffrey
straight or slightly reflexed, thick woody. Apo­physes 731 (holotype E) & USA: California, Trinity Co.,
slightly raised to gibbous, more pronounced on one west side of Scott Mountain, on Trinity – Siskiyou
side of cone towards base, transversely keeled, rhom- County line, R. O. Alava & A. Alava 2439A (epitype
bic to pentagonal in outline, with sometimes crenate E).
upper margins, radially striate, lustrous ochraceous
or light brown. Umbo dorsal, flat to slightly raised, Pinus jeffreyi Balf. var. baja-californica Silba, Phyto-
with a minute, deciduous prickle. Seeds obovoid, logia 68: 52. 1990.
slightly flattened, 3.5–6 × 2–3.5 mm, dark grey-
brown. Seed wings obliquely ovate-oblong, 13–17 ×
Etymology
6–8 mm, yellowish, translucent, with a grey or black
tinge. This species was named after John Jeffrey, a plant
collector who discovered it in 1852 in the Shasta Val-
ley of California.
Distribution
Mexico: Jalisco, in the NW part of the Sierra Madre
Vernacular names
del Sur, on the Pacific slope, mainly in the Sierra de
Cuale (Sierra el Tuito). Jeffrey pine, Jeffrey’s pine
TDWG codes: 79 MXS-JA
Description
Ecology
Trees to 50 m tall, d.b.h. to 1.5 m; trunk mono­podial,
This species occurs at altitudes between 800–1200 straight. Bark thick, rough and scaly, divided by
(–1650) m a.s.l. in mountainous terrain on deep, deep fissures into thick, elongated plates, light buff
acidic soils derived from granitic rocks. The climate to light brown or red-brown, the fissures dark grey-
is subtropical, with a 5–6 month dry season from brown. Branches spreading horizontally or assur-
December to May and annual precipitation from gent at ends, persistent, forming an open pyramidal
1000–1500 mm depending on altitude and exposi- or flat-topped crown. Shoots stout, very rough and
 scaly, light orange-brown, often glaucous. Cata- with an altitudinal range of (50–)300 m to 3050 m
phylls ca. 20 mm long, triangular-subulate, reflexed, a.s.l. It is tolerant of low temperatures in winter and
scarious, with erose-ciliate margins, light brown. can grow on thin soil or even in crevices of bare
Vegetative buds ovoid-acute or subglobose; terminal granite rock. In the Sierra Nevada of California the
bud 15–20 mm long; lateral buds smaller, not resin- species, with its close relative P. ponderosa, is charac-
ous. Fascicle sheaths to 20–25 mm long, persistent teristic of open, dry and summer-warm mixed for-
but reduced to 10–15 mm in older fascicles. Leaves ests of the ‘Yellow pine belt’ where it tends to occupy
in fascicles of 3, sometimes a few fascicles of 2, per- the upper zone towards the tree line. In the south-
sisting (3–)4–5(–6) years, thick, rigid, straight or ern part of this mountain range it occurs in diverse
slightly curved, sometimes twisted, (12–)15–22(–25) mixed coniferous forest with e.g. Pinus ponderosa,
690 cm long, 1.5–1.9(–2) mm wide, with serrulate mar- P. lambertiana, P. monticola, P. contorta, Abies con­
gins, acute-pungent to acuminate, light yellowish color, A. magnifica, Calocedrus decurrens, Junipe­
green to greyish green. Stomata on all faces of leaves, rus occidentalis, and Sequoiadendron giganteum. In
in conspicuous lines. Pollen cones ovoid-oblong southern California and Baja California only Abies
to cylindrical, 2–3.5 cm × 6–7 mm, yellow or pur- concolor, Calocedrus decurrens, Pinus contorta, and
plish yellow. Seed cones subterminal, solitary or in P. lambertiana accompany P. jeffreyi. In the Klamath
pairs on short, stout, persistent peduncles, spread- Mountains of Oregon P. jeffreyi occurs on thin ul-
ing and seemingly sessile at maturity, leaving a few tramafic soils of volcanic origin (peridotites and ser-
basal scales on the branch when falling. Mature pentine) which are poor in nutrients; on these soils
cones broadly ovoid to subglobose, with a slightly its most common associate is Calocedrus decurrens.
oblique, flattened base, (10–)12–25(–30) × 9–14(–16) Here it descends to low elevations (around 100 m),
cm when open. Seed scales parting soon and wide, while in the Sierra Nevada it ascends to 2900 m and
thin woody, straight or slightly curved, up to 20 mm in the Sierra San Pedro Martír of Baja California to
wide. Apophysis slightly raised, transversely keeled, 3050 m.
broadly rhombic in outline, on the proximal scales
more gibbous, often resinous, ochraceous to light
Conservation
brown. Umbo dorsal, moderately raised, transverse-
ly keeled, terminating in a distinct, curved, 3–5 mm IUCN: LC
long spine. Seeds obliquely ovoid, slightly flattened,
9–12 mm long, light yellowish brown with faintly
Uses
darker spots. Seed wings oblique, with a straight and
a curved side, 20–25 × 10 mm, light yellowish brown Jeffrey pine is very similar to Ponderosa pine in its
to light brown. wood properties and is consequently an important
timber tree. It has a ‘preference’ for poorer sites and
higher altitudes, and consequently grows slower
Taxonomic notes
than Ponderosa pine in its natural habitat, but it
Pinus jeffreyi is closely related to P. ponderosa and grows as fast or faster in plantations. The timber in-
P. coulteri and introgression via pollen may occa- dustry does not differentiate the wood from the two
sionally occur where these species occur together. species; details are therefore given under P. pondero­
sa. The resin of P. jeffreyi contains much heptane and
lower levels of terpenes than found in other pines;
Distribution
in the past this biochemical product was distilled
USA: California, W Nevada, S Oregon; Mexico: Baja and used as an additive to raise the octane levels of
California Norte. petrol (gasoline), as well as for medicinal applica-
TDWG codes: 73 ORE 76 CAL NEV 79 MXN-BC tions. Jeffrey pine is not uncommon in cultivation,
but it is mostly restricted to large parks and gardens
with tree collections (arboreta). In forestry, crosses
Ecology
and back-crosses have been experimented with a
Pinus jeffreyi is a montane to subalpine species number of related pines, in particular P. coulteri and
largely confined to the mountains of California, P. ponderosa and its varieties.
Pinus kesiya Royle ex Gordon, Gard. Mag. & Reg. dark brown to nearly black; wing 15–20 mm long,
Rural Domest. Improv. 16: 8. 1840. 6–8 mm wide, persistent.

Etymology Distribution
The species epithet refers to the Khasi Hills in NE China: Yunnan, extreme SE Xizang [Tibet]; NE In-
India, from where it was first reported by J. F. Royle; dia: Assam; Indochina; Malesia: Philippines (Lu-
its spelling has long been a matter of unnecessesary zon).
controversy. Royle’s (European) spelling of the topo- TDWG codes: 36 CHC-YN CHT 40 ASS-AS ASS-MA
nym at the time was as good as any other and must ASS-ME ASS-MI ASS-NA 41 CBD LAO MYA THA VIE
be followed for the name of this pine in botanical 42 PHI 691
nomenclature.
Ecology
Vernacular names
Pinus kesiya occurs in pine savannas, pure stands
Khasia pine, Khasi pine, Luzon pine, Benguet pine; with nearly closed canopy, and mixed pine-broad-
dingsa, far, saral (India); tinyu (Myanmar). leaved forests in valleys with e.g. Quercus serrata
and Alnus nepalensis along streams. It occupies drier
sites in NE India, Myanmar and Thailand, at altitudes
Description
generally between 800 and 1500 m a.s.l., occasional-
Trees to 30–35(–45) m tall; trunk to 1 m d.b.h., often ly to 2000 m. Further east, in Laos, Vietnam and on
straight and columnar. Bark on trunk thick, deeply the island of Luzon in the Philippines, its altitudinal
fissured forming irregular plates, rough and flaking, range is greater and it is found up to 2700–3000 m
brown weathering grey-brown. Branches spread- a.s.l. in a much wetter climate. Despite this, fires are
ing, usually crooked, slowly self-pruning, forming frequent, creating an open, grass-­dominated wood-
a broadly domed crown. Foliage branches slender, land or savanna with scattered stands of pines or
rough with pulvini from fallen leaf fascicles, later solitary trees. The soils are usually sandy or loamy
more or less smooth as these disappear, lustrous yel- and derived from sandstone or quarzite. Pinus kesiya
lowish brown, darkening to orange-brown, shoots is a pioneer in deforested secondary vegetation, es-
multi-nodal within one year. Buds small, conical, pecially if fire has been a factor in the disturbance.
without resin or slightly resinous; cataphylls red-
brown. Leaves in fascicles of 3, held together in a
Uses
15–20 mm long, persistent basal sheath, (10–)12–
22(–25) cm long, slender, very lax or flexible, 0.5– Khasia pine is an important timber tree in much of
0.7(–1) mm wide; margins minutely serrulate; apex SE Asia and has been planted as a forestry tree in
long acuminate; stomata in fine lines on all surfaces. many countries in Africa, South America and Oce-
Pollen cones clustered and spirally arranged at base ania (including Australia). It is a quickly establish-
of new shoots, short cylindrical, yellow. Seed cones ing and fast growing species and as a result its most
solitary or in pairs or threes, short pedunculate, common use is for pulpwood in the paper industry
spreading or reflexed, persistent, 4.5–7(–8) cm long, (class A Kraft paper). In India charcoal burning has
narrowly ovoid-conical when closed, symmetrical been important but has much declined. Other wood
or slightly non-symmetrical, ovoid to subglobose uses are roundwood (poles), construction timber,
when opened, then 3–4(–5.5) cm wide. Seed scales floorboards, plywood, and furniture. It is a tropical
woody, rigid, more or less recurved when spreading, pine, but it does not grow well in the hot and hu-
narrowly oblong, 2.5–3 × 1–1.5 cm, dark brown. Apo- mid lowland tropics; it clearly needs seasonality of
physis prominently raised, more or less pyramidal, precipitation. In the Philippines coffee plantations
irregularly rhombic in outline, transversely keeled, are often established under natural stands of Kha-
light brown. Umbo small, ellipsoid or at least pro- sia pine. The resin is of high quality, but the trees
truding, armed with a very small, persistent prickle. of this species do not yield it freely and therefore
Seeds ellipsoid, 5–7(–8) mm long, slightly appressed, they are not tapped on a large economic scale. In
­horticulture its use is limited to a few arboreta and
 botanic gardens, but in the Philippines it is cultivat- enough to warrant species status and it becomes
ed as a Christmas tree. ­either a synonym of P. kesiya (as spelled by Royle
and Gordon, not by subsequent authors) or of its va-
2 varieties are recognized: riety, first described from Vietnam, but apparently
present in much of the range of the species. The lat-
Pinus kesiya Royle ex Gordon var. kesiya. Pinus ter taxonomy has here been adopted (Farjon, 1998,
insularis Endl. var. khasyana (Griff.) Silba, Phyto- [2001], 2005b), but this remains somewhat tentative
logia 68: 51. 1990. Type: India: Meghalaya, Khasi and other workers on conifers (e.g. Nguyen Tien
Hills, [“regio temp. alt. 2–6000 ft.”], J. D. Hooker & Hiep et al., 2004) have considered it as a synonym of
T. Thomson s.n. (lectotype K). P. ­kesiya if circumscribed slightly wider.
692
Description Distribution
Trees to 35 m tall; trunk to 1 m d.b.h. Leaves long and China: Yunnan; Vietnam; Laos; Thailand; Philip-
lax, 12–25 cm long, 0.5–0.7 mm wide. Seed cones pines.
4.5–7 cm long and 3–4.5 cm wide when opened. TDWG codes: 36 CHC-YN 41 LAO THA VIE 42 PHI

Distribution Conservation
NE India: Assam; China: W Yunnan, extreme SE Xi- IUCN: LC
zang [Tibet]; Indochina: Kampuchea [Cambodia],
Laos, Myanmar [Burma], Thailand.
TDWG codes: 36 CHC-YN CHT 40 ASS-AS ASS-MA Pinus koraiensis Siebold & Zucc., Fl. Japon. 2
ASS-ME ASS-MI ASS-NA 41 CBD LAO MYA THA (3): 28, t. 116. 1842. Type: Illustration in Siebold
& Zuccarini, Fl. Japon. 2: t. 116, f. 5–6. 1842.
(lectotype).
Conservation
IUCN: LC Pinus prokoraiensis Y. T. Zhao, Bull. Bot. Res. North-
East. Forest. Inst. 10 (4): 69. 1990.
Pinus kesiya Royle ex Gordon var. langbianensis
(A. Chev.) Gaussen ex N.-S. Bui, Adansonia, sér. 2,
Etymology
2: 338. 1962. Pinus langbianensis A. Chev., Rev. Int.
Bot. Appl. Agric. Trop. 24: 25. 1944; Pinus insularis The species epithet refers to Korea.
Endl. var. langbianensis (A. Chev.) Silba, Phytologia
68: 51. 1990. Type not designated.
Vernacular names
Pinus insularis Endl., Syn. Conif.: 157. 1847; Pinus Korean pine; channamu (Korea, DPR); hong song
kesiya Royle ex Gordon subsp. insularis (Endl.) (Chinese)
D. Z. Li, Edinburgh J. Bot. 54 (3): 346. 1997.
Description
Description
Trees to 25–30 m tall (in China reported to attain
Trees to 45 m tall. Leaves (10–)12–18 cm long, flex- 50 m); trunk to 1 m d.b.h., growing to a straight,
ible but not lax, 0.7–1 mm wide. Seed cones 5–8 cm columnar bole. Bark smooth in young trees, in
long and 4–5.5 cm wide when opened. larger trees becoming rough and scaly, with shal-
low grooves and longitudinal fissures, forming
oblong, reddish grey or grey-brown plates; inner
Taxonomic notes
bark red-brown. Branches spreading or ascending,
This taxon was apparently first described by Endli- forming a broad crown; top of trees often forked.
cher (1947) as Pinus insularis from the Philippines. Foliage branchlets slender, new shoots occasion-
Later comparisons have found it not to be distinct ally ­pubescent with yellowish hairs, red-brown
­ ecoming grey-brown. Buds oblong or ovoid, the
b with other pines. In Korea and NE China (‘Manchu-
central bud to 10 mm long, slightly resinous; cata- ria’) this pine has been heavily exploited, resulting in
phylls reddish brown. Leaves in fascicles of 5, per- the disappearance of many magnificent pine forests.
sisting 2–3 years, held in a deciduous basal sheath
of orange-brown flimsy scales, slender, straight or
Conservation
abruptly bent, 6–13 cm long, pliant or lax, ca. 1 mm
wide, green on abaxial surface; margins minutely Despite over-exploitation resulting in removal of
serrulate; stomata in bluish white lines on the two many indigenous forests with this species, its wide
adaxial faces. Pollen cones in small clusters at base distribution and partial regeneration (outside af-
of new shoots, short ellipsoid-cylindrical, ca. 15 mm forestation) appears to guarantee the survival of this
long, yellow. Seed cones solitary or in whorls of 2–4 species, which is therefore not considered threat- 693
on stout, 1–1.5 cm long peduncles, semi-persistent, ened under IUCN criteria.
initially erect, becoming spreading to pendulous as IUCN: LC
they grow, blue-green and resinous, when mature
9–14 cm long and 6–8 cm wide, ovoid-oblong, light
Uses
brown or grey-brown. Seed scales at base of cones
reflexed or recurved, more or less indehiscent to Korean pine is a highly important timber tree; in
slightly spreading but not releasing the seeds, more large parts of its range it has been over-exploited,
or less rhombic in outline, the middle scales ca. 3 cm but it is now used widely in afforestation schemes
long, 2.5 cm wide at transition to apophysis, with especially in NE China. Its timber is of good qual-
two deep seed cavities adaxially; apophyses broadly ity, light and soft, straight grained and easy to work
triangular, slightly thickened, oblong at cone apex, with in milling and carpentry. It is fairly decay re-
longitudinally striated or grooved, nearly straight sistant and therefore finds uses like telephone poles,
but recurved or reflexed at the margin towards apex; railway sleepers, wooden bridges, and boat building.
umbo terminal, obtuse or acute. Seeds triangular- In construction it provides building timbers as well
obovoid, 12–16 mm long, light brown or grey-brown, as flooring, plywood and veneers. It can be chipped
without a wing when removed from the scale. for particleboard or flakeboard manufacture, or
pulped for the paper industry. More specialized
uses are furniture, sports equipment and musical in-
Distribution
struments. Resin is extracted from wood pulp and
China: Heilongjiang, Jilin; Japan: Honshu; North & used to produce turpentine and other products. The
South Korea; Russian Far East: Amur, Khabarovsk, seeds are rich in vegetative oil with a high nutritive
Primorye. value, and this is used in the food processing indus-
TDWG codes: 31 AMU KHA PRM 36 CHM-HJ CHM- try; seeds can also be consumed whole and most of
JL 38 JAP-HN KOR-NK KOR-SK the imported pine kernels in Europe and the USA
are now sourced from this species. It is widely culti-
vated as an ornamental tree in China, Korea and Ja-
Ecology
pan, but less common in Europe and the US, where
In the Russian Far East this pine grows from 200 m until recently only a limited number of cultivars was
to 600 m a.s.l., 500 m to 1300 m a.s.l. in China, and in known, but now on the increase.
Japan it occurs to an altitude of 2500 m. The climate
has a summer monsoon character within proxim-
ity of the coast, but with a strong continental influ- Pinus krempfii Lecomte, Bull. Mus. Hist. Nat.
ence further inland. Temperature extremes range (Paris) 27: 191. 1921. Ducampopinus krempfii
from +37 °C to –45 °C within its natural range. Pinus (Lecomte) A. Chev., Rev. Int. Bot. Appl. Agric.
koraiensis grows in dry places on podzols among Trop. 24: 30. 1944. Type: Vietnam: South Vietnam,
deciduous broad-leaved trees like oaks, poplars Tinh Khanh Hoa, near Nha Trang, M. Krempf 1537
and birches, but on the Russian coast of the Sea of (holotype P). Pl. 30
Japan it is codominant with Abies holophylla, form-
ing groves of conifers in a more varied deciduous Pinus krempfii Lecomte var. poilanei Lecomte, Bull.
broad-leaved forest. In Japan it also occurs together Mus. Hist. Nat. (Paris) 30: 325. 1924.
 5

694

1
3

2
4

Pl ate 30. Pinus krempfii. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Seed cone. 5. Seeds.
 should be classified in its own genus (or subgenus)
Etymology
Ducampopinus A. Chev. This opinion was mostly
This species was named after the collector of the based on the peculiar, flat needles; virtually all other
type specimen, M. Krempf. organs, including the dwarf shoots that produce the
needles, are matched by other pines and are mostly
unique to the genus. More recent phylogenetic anal-
Vernacular names
yses based on DNA data have given results that do
Krempf ’s pine; Thong la det (Vietnamese) not support this major distinction and have placed
the species firmly within the genus Pinus and as al-
most certainly belonging to subgenus Strobus (Lis-
Description
ton et al. in Mill, 2003). It is definitely an ancient 695
Trees to 40(–50) m tall; trunk to 3(–4) m d.b.h., erect, relict species, as are several other species in this
straight, massive, often butressed. Bark smooth on ­subgenus.
young trees, becoming rough and scaly on large
trunks, with irregular plates divided by shallow fis-
Distribution
sures, grey-brown to grey. Branches long, spreading
wide and near the top ascending, forming a broadly S Vietnam (southern Truong Son Range, between
domed or flat-topped, dense or more open crown. Dalat and Nhatrang and N of Nhatrang).
Foliage branches slender, multinodal, smooth after TDWG codes: 41 VIE
leaves have fallen, light brown turning grey with
age. Buds small, almost continuously producing new
Ecology
shoots and leaves, not resinous. Leaves in fascicles of
2, with early deciduous basal sheaths consisting of a Pinus krempfii is rare and grows in primary closed
few thin, acute scales which fall separately, spread- evergreen tropical montane rainforest, where it is
ing, straight or slightly curved, 4–5(–7) cm long, an emergent above the general canopy dominated
extremely flat, 1.5–5 mm wide, often twisted in the by angiosperm trees. Its altitudinal range is between
lower, narrower fourth to third of their length, with 1200 m and 2000 m a.s.l. (most common between
a narrow median rib on each side, gradually taper- 1500 m and 1800 m) and it is often found on steep
ing to an acute apex, green on adaxial side, glaucous slopes and on mountain ridges. The mean annual
green on abaxial side; stomata in fine lines on abaxial temperature is 14–20 °C and mean annual rainfall
face. Pollen cones small, in groups at base of new is in excess of 1500 mm, with a dry season from
shoots, 7 mm long when full-grown. Seed cones soli- December to July. It grows predominantly in well-
tary or more commonly in pairs on short peduncles, drained, low-pH lateritic clay soils indicating low
maturing within 2 years, spreading and opening, nutrient availability. Other conifers with which this
falling after seed dispersal, 5–7 cm long, narrowly species is associated are Fokienia hodginsii, Pinus
ovoid when closed, light green, opening to 3–5 cm dalatensis and Dacrydium elatum. It is probably a
wide, becoming brown. Seed scales relatively few, long-lived emergent tree dependent on episodal dis-
soft woody, oblong-concave, dark brown when ma- turbance regimes for successful regeneration, but no
ture; apophyses hard woody, thickened, raised and detailed observations have been made to verify this
often curved, prominently transversely keeled, more and its ecology remains poorly known.
or less rhombic in outline or with rounded upper
margins, light brown or light reddish brown; umbo
Conservation
dorsal, raised, acute or more or less flat with a minute
prickle. Seeds very small, 2.5–3 mm long, more or The recent decline of this species has been attrib-
less ovoid, nearly black; wing long and narrow, 17–20 uted to the effects of the American War (also known
mm long, 4–5 mm wide, detachable from the seed. as Vietnam War) in the 1960s and the clearance of
forested land for agriculture in the following dec-
ades under pressure of rapid population increase. Its
Taxonomic notes
limited distribution and the inaccessibility of much
This remarkable pine has long been considered dis- of its habitat mean that there has been little direct
tinct from all other species, even to the extent that it exploitation. Most populations are now within
 r­ ecently established nature reserves around the Bi oblong; terminal bud 5–10 × 4–6 mm; lateral buds
Doup massif. This species is listed in Group IIA of smaller, not resinous. Fascicle sheaths 10–15 mm
the List of Rare and Precious Flora and Fauna [Viet- long on young fascicles, soon deciduous and absent
nam] for limited use. when the leaves are full-grown. Leaves in fascicles
IUCN: VU (B1+2c) of 5, persisting 2–4 years, straight, slightly twisted,
lax, (3.5–)4–8(–10) cm long, 0.8–1.5 mm wide, with
remotely and sometimes scarcely serrulate margins;
Uses
apex acute-acuminate; leaf colour green to glaucous
This species could be a valuable timber tree be- green. Stomata on all faces of leaves. Pollen cones
cause of its size and good quality of wood, but it is forming more or less elongated ‘spikes’, ellipsoid-­
696 not commercially exploited. Attempts at cultivation cylindric, 10–15 mm long, yellowish. Seed cones at
in Vietnam have remained unsuccessful, due to its ends of main branches, subterminal, solitary or in
apparently very specific conditions for growing. It whorls of 2–4, erect on 5–12 cm long peduncles, be-
is not known to be in cultivation. More serious at- coming pendulous. Mature cones narrowly cylindri-
tempts should be made to grow this pine in areas cal when closed, straight or slightly curved, broadly
with a warm, moist climate, which requires study of cylindrical when opened, then (30–)40–45(–56)
its development from seed to young tree in its natu- × 8–15 cm. Seed scales readily opening, patent or
ral habitat. slightly reflexed, with basal sterile scales often more
reflexed, angular-obovate to obtrullate, up to 40 mm
wide and 4–5 mm thick. Apophysis triangular to
Pinus lambertiana Douglas, Trans. Linn. Soc. obtusely rhombic, 5–8 mm thick at base, straight,
London 16: 500. 1827. Type: USA: Oregon, [locality smooth or radially striated, yellowish brown, often
unknown], D. Douglas s.n. (lectotype K). Pl. 31 with yellow resin blobs. Umbo terminal, obtuse-­
triangular. Seeds obovoid or obliquely obovoid, (10–)
Pinus lambertiana Douglas var. martirensis Silba, 12–15(–18) mm long, 6–10 mm wide, dark brown.
Phytologia 68: 52. 1990. Seed wings broadly obovoid to dolabriform or trun-
cate, 20–30 mm long, 12–15 mm wide, light brown.
Etymology
Distribution
This great pine was named by David Douglas, who
introduced it to England, after Aylmer Bourke Lam- USA: California, Oregon, Nevada (extreme western
bert (1761–1842), a gentleman botanist. part); NW Mexico: Baja California Norte (Sierra
San Pedro Martír).
TDWG codes: 73 ORE 76 CAL NEV 79 MXN-BC
Vernacular names
Sugar pine
Ecology
This majestic pine is common in the mixed and
Description
varied conifer forests of the Transition Zone in the
Trees to 70–75 m tall, d.b.h. to 3–3.8 m; trunk high mountains of Oregon and California common-
straight; bole often free of branches half or more of ly between 600 m and 2400 m a.s.l., but reaching
its length. Bark thick, rough, scaly, on lower part of 2800–3200 m in the south of its range. In the Sierra
trunk breaking into increasingly large, irregular to Nevada of California it is restricted to the western
longitudinal plates separated by deep fissures, cin- slopes at middle elevation, between the warmer Up-
namon to grey-brown or purplish brown. Branch- per Sonoran Zone and the colder Canadian Zone.
es up to 12 m long, spreading horizontally, curved Here one of the most impressive conifer forests in
downwards towards ends; crown broadly conical, in the world occurs, dominated by gigantic trees; in-
old trees often flat-topped from windbreak. Shoots deed the largest tree species in the world, Sequoia­
slender, flexible, puberulent at first but soon gla- dendron giganteum, is restricted to this zone and this
brous, orange-brown turning light grey. Cataphylls mountain range. Other species are Pinus monticola,
small, triangular, scarious. Vegetative buds ovoid- P. ponderosa, P. jeffreyi, P. contorta var. murrayana,
 697

4
1
2

Pl ate 31. Pinus lambertiana. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cone.
 Abies magnifica, A. procera, Calocedrus decurrens, Pinus latteri Mason, J. Asiat. Soc. Bengal. Sci. 18:
and in the north of the range Pseudotsuga menziesii. 74. 1849. Pinus merkusii Jungh. & de Vriese var.
A hospitable climate with warm, sunny summers latteri (Mason) Silba, Phytologia 68: 53. 1990; Pinus
moistened by rain showers and snowy but not ex- merkusii Jungh. & de Vriese subsp. latteri (Mason)
tremely cold winters marks this zone. In Oregon D. Z. Li, Edinburgh J. Bot. 54 (3): 346. 1997. Type
Pseudotsuga menziesii, Abies grandis, Tsuga hetero­ not designated. Fig. 227
phylla, Thuja plicata, and Calocedrus decurrens are
its most common associate conifers. Broad-leaved
Etymology
trees are rare and much smaller, Arbutus menziesii
and Quercus kelloggii are perhaps the most com- This species was named by the Rev. F. Mason after
698 mon. The ground is open, littered with conifer nee- “Capt. Latter, the discoverer”.
dles, interspersed with small, flowery meadows and
lush streamsides full of flowering shrubs. The bright
Vernacular names
yellow lichen Letharia vulpina festoons trunks and
branches everywhere. This type of forest becomes Tenasserim pine; shaja, tinshu (Burmese); son-
resticted to the highest mountain summits further songbai (Thai); nan ya song (Chinese)
south and the diversity of conifers dimishes as does
their size.
Description
Trees to 30 m tall; trunk to 2 m d.b.h. Bark thick
Conservation
and scaly, rough, breaking into numerous small,
IUCN: LC dark grey plates. Branches spreading and assur-
gent, forming a broadly domed or umbrella-shaped
crown in older trees. Foliage branches stout, gla-
Uses
brous, brown or dark brown. Buds cylindrical; ter-
Sugar pine is the most valuable of the pines due to minal bud 1.5–2 cm long, without resin; cataphylls
its enormous size and its light, soft, even-grained, brown. Leaves in fascicles of 2, held by a persistent,
knot-free wood of which very large, straight pieces 15–20 mm long basal sheath, long and slender but
can be sawn. Consequently, old growth stands of rigid, 15–25(–27) cm long, straight, semi-circular
this species command very high prizes; fortunately in transverse section, ca. 1.5 mm wide, dull green;
many are now protected in National Parks and other margins minutely serrulate; apex long acute; sto-
reserves. This species is used for high quality con- mata in numerous lines on all surfaces. Pollen cones
struction timber and the finished milled wood of clustered, spirally arranged, erect, cylindrical, 2–3
this ‘king of pines’ as John Muir called it, makes it cm long. Seed cones sub-lateral on new shoots, usu-
ideal for high standard windows and doors as well as ally solitary, sometimes in pairs, on ca. 1 cm long,
foundry casting and even musical instruments such stout peduncles, spreading at nearly right angles
as organ pipes and piano keys. Plantation forestry from branch, ovoid-conical or oblong-conical when
has not been very successful and in Europe few trees closed, (5–)6–10(–13) cm long, widening to 4–9 cm
have survived for long in arboreta and parks due to and becoming broadly ovoid with a flat base and
white pine blister rust (Cronartium ribicola, Basidio­ spreading scales. Seed scales woody, rigid, oblong,
mycota) to which it is exceptionally sensistive. The twice as long as wide, ca. 3 × 1.2–1.5 cm at mid cone
vernacular name refers to the sugar content of the in the larger cones, light reddish brown; apophyses
resin; in the past native tribes used it as a chewing rhombic or unequally pentagonal in outline, raised
gum. This use was first noted by David Douglas, and strongly transversely keeled, radially striated or
who thought the native people were eating it. ridged, lustrous red-brown to dark brown; umbo
dorsal, more or less central, slightly sunken or flat
to obtuse, unarmed. Seeds ellipsoid to obovoid,
5–8 mm long, 4 mm wide, slightly appressed, grey-
brown, with a 15–20 mm long persistent wing.
 Taxonomic notes Conservation
The occurrence of a so-called grass stage in the seed- IUCN: NT
lings of Pinus latteri and its close relative P. merkusii
has been variously reported. Cooling & Gaussen
Uses
(1970) first described a grass stage in P. merkusii
(Sumatera, Philippines), which was said to be ab- Tenasserim pine is a timber tree used in SE Asia for
sent in “Pinus merkusiana” (an invalidly published light construction purposes. The wood is moder-
name = P. latteri) from mainland Asia. This observa- ately hard and very resinous and only suitable for
tion was reiterated in the second edition of my book indoor applications when processed as sawn timber.
Pines (Farjon, 2005b), but Vidaković (1991) stated Doors, window frames and flooring are common 699
that the grass stage is absent in trees from Sumatera uses; some provenances can be put to veneers. Large
(P. merkusii), but present in trees from the mainland quantities are nowadays used in the wood pulp in-
(P. latteri). Information in PROSEA, a database on dustry, while charcoal burning is a more traditional
plant resources of SE Asia (PROSEA, 1993) likewise use still current. It is also an important tree for resin
mentions the grass stage for “mainland P. merkusii” tapping in some countries. In China, the resin of
= P. latteri. If this information is authentic, and I see this species has some medicinal applications, e.g.
no reason to doubt it, then perhaps this distinction in making ointments. This tropical pine has been
between the two species does not hold true. In that introduced for afforestation in several countries in
case, it could be argued that the other morphologi- Africa, but the results have on the whole been un-
cal differences between P. latteri and P. merkusii are satisfactory because of difficulties with raising it
minor and mostly of a quantitative nature, character beyond the seedling stage in all situations where
states that may only justify distinction as a variety or competition from other plants (‘weeds’) is a factor.
subspecies for P. latteri under P. merkusii, the earlier
name at species rank.
Pinus lawsonii Roezl ex Gordon, Pinetum Suppl.:
64. 1862 [“lawsoni”]. Type: Mexico: Michoacan,
Distribution
Uruapán, Cerro de la Cruz, C. G. Pringle 10141
China: Guangdong (Hainan Island, introduced?), (neotype NY).
Guangxi; Indochina: SE Myanmar [Burma], Kam-
puchea [Cambodia], Laos, Thailand, Vietnam. Pinus lawsonii Roezl ex Gordon var. gracilis Debre-
TDWG codes: 36 CHH(?) CHS-GX 41 CBD LAO MYA czy & Rácz, Phytologia 78 (3): 19. 1995.
THA VIE
Etymology
Ecology
Roezl named this species after the English gardener
Pinus latteri is a widespread species occurring from and plantsman Charles Lawson (1794–1873), who
near sea level to ca. 1200 m a.s.l., forming more or validated David Douglas’ name Pinus ponderosa in
less open stands on old river terraces with sandy the same way as did George Gordon with this name
or gravelly soil or in seasonally dry hills. It is a fire- from him.
adapted species capable of surviving grass fires
through a ‘grass stage’ as a seedling (see also under
Vernacular names
Taxonomic Notes) and can quickly invade open ter-
rain especially in nutrient-poor soils. Its range is Lawson’s pine
within the influence of SE monsoons, with mean
annual precipitation around 1500 mm in Myanmar
Description
and Thailand. It is also a constuent tree in diptero-
carp forests (e.g. Dipterocarpus tuberculatus), espe- Trees to 25–30 m, d.b.h. to 75 cm; trunk mono­
cially in more open situations on drier sites of ridges podial, straight, sometimes tortuous. Bark thick,
and spurs in the hill country of Myanmar [Burma] rough and scaly, with broad and deep longitudinal
and other countries in mainland SE Asia. fissures; outer bark dark blackish brown. Branches
 spreading horizontally, assurgent in the higher part
Conservation
of the crown; forming a broadly domed but open
and irregular crown. Shoots smooth, orange-brown, IUCN: LC
often glaucous. Cataphylls subulate-lanceolate, ca.
10 mm long, scarious, erose-ciliate at margins, dull
Uses
brown. Vegetative buds ovoid-oblong to cylindrical;
terminal bud 10–15 mm long; lateral buds shorter, Along with other pines, which are often more abun-
acute, not resinous. Fascicle sheaths of young leaves dant in the mixed pine or pine-oak forests, this spe-
up to 25 mm long, persistent but reduced to 10–15 cies is cut commercially for lumber, although most
mm in older fascicles. Leaves in fascicles of 3–4 (–5), trees are of medium size and the boles sometimes
700 very seldom 2, persisting 2–3 years, straight or near- tortuous. In several places with abundant trees of
ly straight, rigid, 12–20(–25) cm long, 1.0–1.2 (–1.5) this species, resin may be tapped as well. As an orna-
mm wide, serrulate at margins, acute-pungent, mental tree this species will only grow successfully
glaucous green. Stomata on all faces of the leaves. in countries with very mild climate; it is not known
Pollen cones ovoid-oblong to cylindrical, 1–2 cm × to be in cultivation for this purpose.
5–6 mm, yellowish green, turning light brown. Seed
cones subterminal, sometimes solitary, commonly
opposite on short, stout, curved peduncles which Pinus leiophylla Schiede ex Schltdl. & Cham.,
usually fall with cone. Mature cones narrowly ovoid Linnaea 6: 354. 1831.
to ovoid-attenuate when closed, more or less asym-
metrical, ovoid when open, with a more or less flat-
Etymology
tened base, then 5–8(–9) × 4–6(–7) cm. Seed scales
thick woody, oblong, recurved when fully spread. The species epithet is derived from Latin: lenis = soft
Apophysis slightly raised, transversely keeled, rhom- and Greek: phyllos = leaf.
bic in outline, with undulate-crenate upper margin,
radially striate, light brown to grey-brown. Umbo
Vernacular names
dorsal, pyramidal, curved, lacking a distinct prickle.
Seeds obovoid, slightly flattened, 4–5 mm long, dark Smooth-leaved pine; Chihuahua pine; pino chino
brown. Seed wings 12–16 × 5–6 mm, translucent, (Mexico)
light brown with a dark tinge.
Description
Distribution
Trees to 15–30(–35) m tall, d.b.h. to 20–85 cm; trunk
Mexico: Michoacán, México, Morelos, Distrito Fed- monopodial, erect. Bark very thick, rough, scaly,
eral, (one locality in) Veracruz, Guerrero and Oax- divided into elongated, irregular, scaly plates and
aca. deep, mainly longitudinal fissures, dark grey-brown.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXG- Branches long, slender, spreading or ascending, but
VC MXS-GR MXS-MI MXS-OA lower ones curved down, forming an open, rounded
crown. Shoots more or less scaly, initially reddish
brown or glaucous, soon grey-brown. Cataphylls
Ecology
ca. 6 mm long, lanceolate-subulate, soon recurved,
Pinus lawsonii has an altitudinal range of 1300–2600 light orange-brown. Vegetative buds ovoid-conical,
m a.s.l. It is a constituent of warm-temperate to tem- obtuse; terminal bud 10–15 × 5–7 mm; lateral buds
perate montane forest or woodland, with annual smaller, not or slightly resinous. Fascicle sheaths 12–
precipitation ranging from 600–1500 mm and a dry 20 mm long, early deciduous leaving the full-grown
season from November to May. It is most commonly fascicles devoid of a sheath. Leaves in fascicles of
found in pine-oak forest, also in pine forest with e.g. (2–)3–5(–6), spreading in lax or rigid tufts near ends
Pinus pringlei, P. patula, P. montezumae, P. oocarpa, of branches, persisting 2–3 years, sometimes sub-
P. leiophylla, P. herrerae, P. teocote, and P. pseudo­ pendulous, straight, variable in length on the same
strobus. On sites with sandy, shallow soil Juniperus branch, (4–)6–15(–17) cm long, 0.5–1.3(–1.5) mm
can be codominant. wide; margins (minutely) serrulate; apex acute or
acute-pungent; leaf colour light green or glaucous and snow are common in winter. It is one of the few
green. Stomata on all faces of leaves. Pollen cones pines with a capacity to coppice. The seed cones
cylindrical, 1–1.5(–2) cm × 4–6 mm, yellowish pink. take three seasons to mature, which is exceptional
Seed cones subterminal, solitary or in whorls of 2–5 in pines. Although largely sympatric with var. leio­
on stout, 10–20 mm long, spreading or recurved phylla, the altitudinal range of var. chihuahuana is
peduncles, persisting a few years after seed disper- narrower: 1500–2700(–2950) m, but this is in part
sal. Mature cones (narrowly) ovoid when closed, due to the fact that it does not occur on some of the
nearly symmetrical, (broadly) ovoid when opened, high volcanos of central Mexico. In the Sierra Ma-
(4–)5–7(–8) × (3–)4–5.5 cm when open. Seed scales dre Occidental its lower limit usually is below the
straight or recurved near base of cone, oblong, with level of var. leiophylla. Here it may be found in semi-
nearly straight margins. Apophysis raised, trans- arid habitat with P. cembroides, Juniperus spp. and 701
versely keeled, with the central section of the second xerophytes like Opuntia and Arctostaphylos. Higher
season’s growth distinctly marked as a narrow band up, its habitat is similar to that described above. Its
around the umbo, rhombic in outline, dull brown to poor growth in some areas is related to lower pre-
grey-brown. Umbo dorsal, pyramidal, with a blunt cipitation and rocky, poor, shallow soils especially at
prickle, darker than the apophysis. Seeds obliquely lower and middle altitudes.
ovoid, slightly flattened, 3–4(–5) mm long, dark
grey-brown with black spots. Seed wings obliquely
Uses
oval, 10–18 × 4–8 mm, yellowish brown, translucent,
with a dark tinge. Along with many other species of pine in the Sierra
Madre Occidental, Pinus leiophylla has been heav-
ily exploited for timber in the latter half of the 20th
Distribution
century. Due to its high resin content it does not
SW USA: SE Arizona, SW New Mexico; in Mexico provide high quality wood but at the same time is a
along the Sierra Madre Occidental, the “Eje Volcáni- producer of good resin, for which it is tapped. The
co Transversal” and in the Sierra Madre del Sur as wood is used in heavy construction and for crates
far SE as the highlands of central Oaxaca. and boxes. The variety chihuahuana is less exploited
TDWG codes: 76 ARI 77 NWM 79 MXC-DF MXC- for timber, but an equally prolific producer of resin.
ME MXC-MO MXC-PU MXC-TL MXE-AG MXE-CU Smooth-leaved pine has been planted in various
MXE-DU MXE-GU MXE-HI MXE-QU MXE-SL MXE- parts of the world, especially in Africa, as a planta-
ZA MXG-VC MXN-SI MXN-SO MXS-CL MXS-GR tion forestry tree.
MXS-JA MXS-MI MXS-NA MXS-OA
2 varieties are recognized:
Ecology
Pinus leiophylla Schiede ex Schltdl. & Cham. var.
Pinus leiophylla var. leiophylla is a widespread con- leiophylla. Type: Mexico: Veracruz, Cruz Blanca,
stituent of montane to high montane pine and pine- [“inter Cruz Blanca et Jalacingo”], C. J. W. Schiede
oak forests on deep, well drained soils derived from 1109 (holotype HAL).
various, but usually volcanic or metamorphic rock.
Its altitudinal range is (1500–)1900–2900(–3300) Pinus lumholtzii B. L. Rob & Fernald var. microphylla
m a.s.l., gradually increasing from north to south. Carvajal, Phytologia 59: 135. 1986.
It occurs most frequently with Quercus spp. and/or
with Pinus patula, P. pringlei, P. teocote, P. lawsonii,
Description
P. pseudostrobus, P. montezumae, P. douglasiana,
P. durangensis, and at lower altitudes P. oocarpa. In Trees to 20–30(–35) m tall, d.b.h. to 50–85 cm. Shoots
the NW of its range P. arizonica, P. engelmannii and reddish brown, sometimes glaucous. Leaves in fasci-
P. leiophylla var. chihuahuana grow commonly with cles of (4–)5(–6), 4 more often than 6, (6–)8–15(–17)
it. Locally, Juniperus spp. or Cupressus lusitanica are cm long, 0.5–0.9 mm wide. Stomata in (3–)4–6(–7)
found with it. Annual precipitation varies greatly lines on the convex abaxial face, in (2–)3–4 lines on
with locality and altitude, from a low of ca. 700 mm each adaxial face. Resin ducts in the leaves (1–)2–3
to 1950 mm. In the north and at high altitudes, frost (–4), medial, occasionally 1 internal.
 Pinus longaeva D. K. Bailey, Ann. Missouri Bot.
Distribution
Gard. 57: 243. 1971. Pinus aristata Engelm. var. lon­
Mexico: in NE Sonora, W Chihuahua, Durango, Na- gaeva (D. K. Bailey) Little, Phytologia 42: 221. 1979;
yarit, Zacatecas, Jalisco, Michoacán, México, D. F., Pinus balfouriana Balf. subsp. longaeva (D. K. Bai-
Hidalgo, Morelos, Tlaxcala, Puebla, Veracruz, Guer- ley) E. Murray, Kalmia 12: 23. 1982; Pinus aristata
rero and Oaxaca. Engelm. subsp. longaeva (D. K. Bailey) E. Murray,
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC- Kalmia 13: 21. 1983. Type: USA: Nevada, Wheeler
PU MXC-TL MXE-CU MXE-DU MXE-HI MXG-VC Peak Scenic Area, Wheeler Peak, D. K. Bailey &
MXN-SO MXS-GR MXS-JA MXS-MI MXS-NA MXS-OA J. E. Whitson 7001 (holotype COLO). Fig. 228

702
Conservation Etymology
IUCN: LC The species epithet longaeva refers to the longevity
of this pine.
Pinus leiophylla Schiede ex Schltdl. & Cham. var.
chihuahuana (Engelm.) Shaw, [Pines Mexico]
Vernacular names
Publ. Arnold Arbor. 1: 14. 1909. Pinus chihuahuana
Engelm., in Wislizenus, Mem. Tour N. Mexico: Great Basin Bristlecone pine, Intermountain Bris-
103. 1848; Pinus leiophylla Schiede ex Schltdl. & tlecone pine
Cham. subsp. chihuahuana (Engelm.) E. Murray,
Kalmia 12: 23. 1982. Type: Mexico: Chihuahua,
Description
Cosiquiriachi, F. A. Wislizenus 232 (holotype MO).
Trees, small to medium, height to 16 m; trunk d.b.h.
to 3 m. Trunk monopodial, straight or contorted, of-
Description
ten strongly tapering in old trees or sometimes mul-
Trees to 15–25 m tall, often not more than 10 m, d.b.h. ti-stemmed, branching very low and much reduced
to 20–60 cm. Shoots reddish brown, often glaucous. in size at tree line. Bark thin, shallowly to deeply fis-
Leaves in fascicles of (2–)3(–4, rarely 5), (4–)6–12(– sured, breaking into irregular, scaly ridges, reddish
14) cm long, 0.9–1.3(–1.5) mm wide. Stomata in brown, weathering grey. Branches of first order con-
(4–)5–8(–9) lines on the convex abaxial face, in 3–4 torted, ascending or spreading, persistent, of higher
lines on each adaxial face. Resin ducts in the leaves orders flexible and often pendent. Shoots puberu-
(3–)4–6(–7), medial, occasionally 1–2 internal. lent, with short, non-decurrent pulvini, pale red-
brown, ageing to yellowish grey or grey. Cataphylls
5–7 mm long, subulate, scarious, brown, with entire
Distribution
margins, soon deciduous. Vegetative buds ovoid and
USA: SE Arizona, SW New Mexico; Mexico: along acute, resinous; terminal bud 8–10 mm long; lateral
the Sierra Madre Occidental in NE Sonora, W Chi- buds smaller, all with imbricate, appressed, subulate,
huahua, Durango, Nayarit, N Jalisco and Zacatecas. red-brown scales. Fascicle sheaths deciduous and
TDWG codes: 76 ARI 77 NWM 79 MXE-CU MXE- absent in second year’s fascicles. Leaves in fascicles
DU MXE-ZA MXN-SO MXS-JA MXS-NA of 5, in dense tufts persisting up to 30 years, mostly
connivent, curved forward against shoot, (1.5–)2–
3.5(–4) cm long, 0.8–1.2 mm wide, lacking resin
Conservation
droplets or only a few present; margins entire or dis-
IUCN: LC tally serrulate; apex acute or short acuminate; leaf
colour deep green on the smooth abaxial face, white
stomatal lines on both adaxial faces. Pollen cones
crowded at the proximal end of a new shoot, spread-
ing, subtended by light brown bracts, ellipsoid or
ovoid, ca. 10 mm long, red or purple (sometimes yel-
low), maturing to brown. Microsporophylls peltate,
the distal part cordate to rounded, smooth, ca. 1 mm
wide. Seed cones subterminal, solitary or in pairs, is unlikely that this species would compete success-
nearly sessile, falling after seed dispersal. Immature fully, given its very slow growth. Virtually all popu-
cones erect, ovoid, dark purple (sometimes green), lations now fall under some form of legal protection;
maturing in two seasons. Mature cones ovoid-cylin- many are in reserves or protected wilderness areas
dric becoming more ovoid when open, 6–10 × 4–6 within National Forest or other government owned
cm. Apophyses on seed scales thick, triangular to land. There is not much that active management of
rhombic, transversely keeled, recurved, red-brown. the environment could reasonably do to increase
Umbo dorsal, transverse-triangular at base, termi- survival of seedlings within its natural habitat.
nating in a slender but rigid, variable prickle 1–6 IUCN: VU (B1+2e)
mm long. Seeds ellipsoid or obovoid, 5–8 × 3–5 mm,
light brown with darker red-brown specks. Seed 703
Uses
wings 10–13 mm long.
Great Basin bristlecone pine is now a protected tree
in its native habitat. It was never a timber tree due
Distribution
to extremely slow growth and trunks of any usable
USA: E California, Nevada, Utah. size being contorted or otherwise ‘mis-shaped’ from
TDWG codes: 76 CAL NEV UTA a lumberman’s viewpoint. Dead wood lends itself
beautifully for wood turning and may have been
used before the gathering of this commodity was
Ecology
outlawed. This tree has become famous in the sci-
Like its relatives Pinus aristata and P. balfouriana, ence of dendrochronology, which was pioneered by
P. longaeva is a subalpine to alpine species and like E. Schulman in the 1960s. Ages of individual living
the others a xerophyte, adapted to extreme condi- trees exceeding 4000 years were found, with some
tions and fluctuations of temperature and moisture. between 4500 and 5000 years. Analyzing tree ring
Its altitudinal range is between 1700 and 3400 m patterns of these trees and adding those of dead
above sea level and it often grows with P. flexilis, but trees (which in the dry climate at high altitude do
with few other trees or shrubs, and often in bare, not rot) a tree ring calendar of 8200 years has been
rocky ground. Weather conditions are often extreme constructed, enabling archaeologists to date their
in all seasons, ranging from heat, extreme radiation finds precisely if good pieces of wood are preserved.
and desiccating winds to snow and ice storms that It even led to the ‘calibration’ of dates based on C14
blast the bark off the trunks. Growth is therefore (14C) isotope decay and the realization that the rate
exceedingly slow, producing very dense and hard of this decay has not been entirely constant. This
wood with annual increments of minimal width, of- species is not common in cultivation; it is a relatively
ten restricted to only parts of the trunk and some recent taxonomic split from P. aristata, which can be
of the branches, the remainder being already dead found more often in collections or in large rock gar-
wood. Yet these trees even when 4000 years old still dens. Its very slow growth under conditions of stress
produce cones with viable seeds, and seedlings and should make it an ideal plant for bonsai culture.
saplings can be found growing around them.

Pinus luchuensis Mayr, Bot. Centralbl. 58 (5): 149.

Conservation
1894. Type: Japan: Ryukyu Islands, H. Mayr s.n.,
Pinus longaeva is considered vulnerable to extinc- March 1891 (type not located, A?, PC?).
tion because of its fragmented occurrence in a lim-
ited number of populations, often with only a few
Etymology
hundred or at most 500–1000 mature individual
trees. Recruitment is very slow and it is uncertain, This species is named after the Luchu Islands [Ry-
given the great age of these trees, whether this con- ukyu Islands] in the East China Sea.
stitutes sufficient replacement of (eventually) dying
old trees. It is also uncertain what effects climate
Vernacular names
change may have, if for instance conditions became
more suitable for other conifers at these locations it Luchu pine, Okinawa pine
 and shrubs stabilizing drifting sand, virtually free
Description
from competition by other trees.
Trees to 15(–20) m tall; trunk to 60 cm d.b.h. Bark on
trunk rough and scaly, breaking into large plates on
Conservation
lower part of trunk, but thinner, smooth and greyish
on young trees and on branches in crown. First or- IUCN: LC
der branches few, long and spreading (trees in natu-
ral habitat are mostly wind-deformed), higher order
Uses
branches assurgent and densely crowded, forming
flat-topped or domed crowns in natural habitat. This pine is not important as a timber resource and
704 Foliage branches glabrous, more or less smooth, as such only of local use. It is well adapted to salt-
grey. Buds ovoid-conical, 10–15 mm long, 5–7 mm laden sea winds and has been planted to stablize
wide, resinous; cataphylls appressed, orange or rusty sand dunes in coastal areas in other parts of Japan
brown. Leaves in fascicles of 2, held by a persist- as well as in Taiwan. It is not known to be used as an
ent, slender basal sheath ca. 10 mm long, straight ornamental tree and no cultivars are known of this
or slightly curved, 10–15 cm long, slender, pliant, species.
slightly twisted, 0.7–1 mm wide; margins minutely
serrulate; apex acute; stomata in fine lines on all
surfaces. Pollen cones clustered, spirally arranged, Pinus lumholtzii B. L. Rob & Fernald, Proc. Amer.
short cylindrical, 1.5–2 cm long, yellow tinged with Acad. Arts 30: 122. 1894. Type: Mexico: Chihuahua,
red, becoming reddish brown. Seed cones soli- Coloradas, C. V. Hartman 541 (holotype GH).
tary or sometimes in pairs, persistent on short pe-
duncles, spreading, 4–5.5 cm long, narrowly ovoid
Etymology
when closed, widening to 2.5–3 cm when open. Seed
scales thin woody, rigid, oblong, ca, 2 × 1 cm at mid This species was named after the plant collector Carl
cone, straight or slightly recurved when spreading, Lumholtz (fl. 1900).
dull brown; apophyses rhombic in outline or with
rounded upper margin, slightly raised, transverse-
Vernacular names
ly keeled, lustrous brown; umbo small, pyramidal,
armed with a sharp, mucronate prickle. Seeds ellip- Lumholtz’s pine; pino triste (Spanish)
soid-ovoid, 4–5 mm long, slightly flattened; wing
10–15 mm long, persistent.
Description
Trees to 20 m tall, d.b.h. to 50–70 cm; trunk mono­
Distribution
podial, straight. Bark thick, scaly, very rough, divid-
Japan: Ryukyu Islands (Okinawa, Amami-shoto). ed into elongated, irregular plates and deep, wide,
TDWG codes: 38 NNS longitudinal fissures, greyish brown to dark grey.
Branches spreading or ascending, branches of high-
er orders slender, flexible, drooping. Crown broad,
Ecology
rounded and usually open. Shoots initially glau-
Pinus luchuensis is a maritime pine, occurring in cous, turning reddish brown, then grey. Cataphylls
coastal areas of Okinawa and on smaller islands subulate, up to 10 mm long, scarious, red-brown.
from near sea level to 700 m a.s.l. It grows on sand Vegetative buds ovoid-conical, acute; terminal bud
dunes as well as on rocky outcrops and hillsides, of- ca. 15 × 8 mm; lateral buds smaller, resinous. Fasci-
ten on very wind-exposed slopes or close to the sea, cle sheaths (20–)25–35 mm long on juvenile fasci-
where it becomes wind distorted. This species is very cles, consisting of ca. 10 reddish brown scales, soon
tolerant of saline air and ocean spray, but grows well coming apart, forming a tuft of curling scales at the
in planted situations away from strong winds and base of fascicles before they fall at maturity, leaving
airborne salt. In its natural habitat it mostly forms fascicles bare. Leaves in fascicles of 3 (exception-
pure, open stands with a low vegetation of grasses ally 2 or 4), usually extremely pendulous, persisting
two years, lax but not thin, (15–)20–30(–40+) cm
long, (1–)1.2–1.5 mm wide, with serrulate margins,
Uses
acute, light green. Stomata on all faces of leaves, in
conspicuous lines. Pollen cones sparsely clustered, Pinus lumholtzii is generally known as ‘pino triste’
cylindrical, 2–3 cm × 5 mm when mature, at first for its striking pendulous foliage. It has been over-
pinkish, turning yellow. Seed cones subterminal or exploited with other pines in some areas for timber,
lateral, solitary (sometimes in whorls of 2, rarely 3) but due to its scattered occurrence, especially in
on 10–15 mm long, curved peduncles, breaking off pine-oak forest and in drier sites, it is not a commer-
easily and remaining with the soon deciduous cones. cially important tree. This species, despite its strik-
Mature cones ovoid to ovoid-attenuate when closed, ing foliage, is not known in cultivation.
ovoid to ovoid-acute when opened, then (3–)3.5–
5.5(–7) × (2.5–)3–4.5 cm. Seed scales thick woody, 705
oblong, spreading wide or lower scales remaining Pinus luzmariae Pérez de la Rosa, Bol. Inst. Bot.
closed, with smaller basal scales remaining con- (Guadalajara) 5 (1–3): 127. 1998. Pinus oocarpa
nate. Apophyses slightly raised, thickened along the Schiede ex Schltdl. f. trifoliata Martínez, Anales
distal margin, obscurely transversely keeled, those Inst. Biol. Univ. Nac. México 16: 297. 1945; Pinus
on basal scales gibbous, rhombic to pentagonal in oocarpa Schiede ex Schltdl. var. trifoliata Martínez,
outline, ochraceous to reddish brown. Umbo dor- Pinos Mexic., ed. 2: 308, f. 252–253. 1948. Type:
sal, often tilted towards distal margin of apophysis, Mexico: Durango, Sierra de Chavarria, M. Martínez
flat or slightly raised, with a minute, soon deciduous 3458 (holotype MEXU).
prickle. Seeds obliquely obovoid, slightly flattened,
3–5 mm long, dark brown, often with black dots.
Etymology
Seed wings oblique, widest below apex, (8–)10–14 ×
4–6 mm, light yellowish or greyish brown. This species was named after it’s author’s village
school mistress Luz Maria Villareal de Puga, who
first taught him about plants.
Distribution
Mexico: Sierra Madre Occidental, in Chihuahua, Si-
Vernacular names
naloa, Durango, Nayarit, Jalisco, Zacatecas, Aguas-
calientes and Guanajuato. No common names have been recorded for this little
TDWG codes: 79 MXE-AG MXE-CU MXE-DU MXE- known species.
GU MXE-ZA MXN-SI MXS-JA MXS-NA
Description
Ecology
Trees to 10–15 m tall, d.b.h. to 40–60 cm. Leaves in
The altitudinal range of this species is (1500–)1700– fascicles of 3, rarely of 4, straight, rigid, (11–)14–17
2600(–2900) m a.s.l., which corresponds to the low- (–20) cm long, 1.2–1.6 mm wide. Seed cones 3–5.5 ×
er and middle slopes of the Sierra Madre. It grows 3–5(–6) cm when open.
usually mixed with several species of Quercus and
other pines in pine-oak forest, or on the wetter west-
Taxonomic notes
ern slopes of the Sierra Madre, in mixed pine forest.
Associated pines are e.g. P. leiophylla, P. arizonica, This taxon was until recently known as Pinus oocar­
P. douglasiana, P. teocote, and P. oocarpa; on more pa var. trifoliata (Farjon & Styles, 1997), but Pérez de
mesic sites one can expect P. ayacahuite and Pseu­ la Rosa (op. cit.) has made a case for its recognition
dotsuga menziesii, while in dry habitats Pinus cem­ as a species, finding additional characters besides
broides can occur with it. The annual precipitation, the numbers of needles in a fascicle to distinguish
mostly as summer rains, is a moderate 500–600 mm it. The scattered distribution accross a large part of
(except on the driest and wettest sites). Mexico and beyond, always within the very wide
range of P. oocarpa, and the rarity of the trees with
these characters in any location, seem to go against
Conservation
this, but it can be given the benefit of the doubt in
IUCN: LC
 the absence of evidence, e.g. based on DNA data, to yellowish brown, sometimes glaucous, turning light
the contrary. brown in the second or third year. Vegetative buds
small, ovoid-cylindric; apical bud ca. 10 mm long,
non-resinous; cataphylls brown. Leaves in fasci-
Distribution
cles of 2, sometimes of 3, held in persitent fascicle
Mexico; Honduras (distribution imperfectly sheaths 15–20 mm long, spreading wide, 12–20 cm
known). long, straight or slightly contorted, slender, pliant,
TDWG codes: 79 MXE-DU MXS-JA MXS-MI MXS- ca. 1 mm wide, slightly twisted; margins minutely
OA 80 HON serrulate; leaf colour bright green; stomata in fine
lines on all surfaces. Pollen cones in clusters near
706 base of new shoots, spirally arranged, short cylindri-
Ecology
cal, yellow to orange-brown. Seed cones mostly soli-
Similar to that of Pinus oocarpa. tary on short peduncles, spreading or down-curved,
falling soon after seed dispersal, variable in size,
3–9 cm long, narrowly ovoid when closed, ovoid
Conservation
when opened and then 2.5–5 cm wide, symmetrical
IUCN: DD or nearly so. Seed scales thin woody, not spreading
widely, oblong to obovoid-oblong; apophyses nearly
flat or slightly raised, sub-rhombic in outline, with
Uses
rounded upper margin and cuneate lower margin,
No uses have been recorded, but it will be used in transversely keeled in a down-curved line, becom-
the same way as Pinus oocarpa. ing chestnut-brown; umbo dorsal, more or less cen-
tral, obtuse or shortly mucronate. Seeds more or less
obovoid, 4–8 mm long, dark brown; wing 12–21 mm
Pinus massoniana Lamb., Descr. Pinus 1: 17, t. 12. long, 5–7 mm wide, persistent.
1803.
Distribution
Etymology
Central and SE China (accross 15 provinces); Tai-
This species commemorates the botanist Francis wan.
Masson (1741–1805) who never visited China and is TDWG codes: 36 CHC-CQ CHC-GZ CHC-GZ CHC-
best known for his plant collecting at the Cape of HU CHC-SC CHH CHN-SA CHS-AH CHS-FJ CHS-GD
Good Hope in South Africa in the 1770s. CHS-GX CHS-HE CHS-HN CHS-JX CHS-ZJ 38 TAI

Vernacular names Ecology

Masson’s pine, Chinese red pine; ma wei song (Chi- Pinus massoniana is a pine which grows as well in
nese) the lowlands as in the highlands, and occurs from
a few meters to an altitude of about 2000 m a.s.l.
Because of this, it occurs in greatly varying clima-
Description
tological conditions, from the moist lowland river
Trees to 45 m tall; trunk to 1.5 m d.b.h., usually a valleys to the dry mountain plateaus of the interior
straight bole. Bark in the upper part of stem and parts of China. In general, it is a conifer inhabiting
crown thin, flaking, reddish brown; lower down a region with a warm temperate climate. Extensive
trunk becoming thick, with long vertical fissures, stands of this pine occur in several areas where cul-
rough and scaly, grey-brown or red-brown. Branch- tivation of the land has been marginal, i.e. in hilly
es spreading and ascending, self-pruning, higher or rocky terrains and on the poorer acidic soils
order branches more or less pendulous, forming a of mountain slopes and ridges. This species is not
rounded or domed crown. Foliage branches slender, found in limestone formations. It occurs often in
rough with decurrent pulvini after leaf fascicles have mixed light forest or woodland with angiosperms
fallen; shoots elongating twice per year, glabrous, such as Lithocarpus and Quercus (Fagaceae) and
some other conifers, e.g. Keteleeria davidiana and
Distribution
Cunninghamia lanceolata. It is often a pioneer spe-
cies invading abandoned agricultural fields or sec- South Central and SE China; Taiwan.
ondary vegetation in forest clearings. TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
SC CHN-SA CHS-AH CHS-FJ CHS-GX CHS-HN CHS-
JX 38 TAI
Uses
Masson’s pine is one of the most important conifer
Conservation
trees in China. It is fast growing and widely used in
plantations grown for wood, pulp, resin and other IUCN: LC
products. The wood is of good quality and used 707
for construction as well as the pulp and wood fibre Pinus massoniana Lamb. var. hainanensis
industries. Natural stands, especially old growth, W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4):
provide most of the sawn timber, while fast grow- 85. 1975. Type: China: Hainan, Wang Chin 3117
ing plantations are mostly converted into pulpwood. (holotype PE).
Many uses are given to the sawn timber, ranging
from poles, railway sleepers and heavy or light con-
Description
struction and plywood to boxes, furniture, veneers,
cooperage, tool handles, wood carving, and musical Bark red-brown towards base of trunk. Seed cones
instruments. Pine resin of this species forms the basis ovoid when opened, 4–7 × 2.5–5 cm.
for a variety of chemical products, the so-called na-
val stores industry. China is now the world’s leading
Taxonomic notes
producer of gum rosin, both for domestic consump-
tion and export, and Masson’s pine is the major pro- In Flora of China 4 (1999) two varieties are recog-
vider of this, supplying 95% of China’s oleoresin and nized besides var. massoniana; one of these, var.
employing 200,000 workers (Langenheim, 2003). shaxianensis D. X. Zhou is in Farjon (2005b) con-
Oil is extracted from the needles, and this and other sidered to be synonymous with var. massoniana.
parts of the tree are extensively used in traditional The other variety, var. hainanensis W. C. Cheng &
Chinese medicine. Edible mushrooms are cultivated L. K. Fu, was described from Hainan Island in south-
on bedlogs of this pine laid out in limestone caves. ern China and has red-brown bark not only in the
Masson’s pine is not widely planted outside China, upper parts of the tree, but also on the lower trunk.
and is rather uncommon as an amenity tree in re- The cones seem to be less variable in size and more
gions with a mild climate. consistently ovoid and perhaps smaller, but there
may be much overlap of these continuous measure-
2 varieties are recognized: ments with cones of var. massoniana on the main-
land. This variety is here only tentatively accepted.
Pinus massoniana Lamb. var. massoniana. Type:
Illustration in Lambert, Descr. Pinus 1: t. 12. 1803
Distribution
(lectotype).
China: Hainan Island (Yajiadaling).
Pinus crassicorticea Y. C. Zhong & K. X. Huang, Gui- TDWG codes: 36 CHH
haia 10 (4): 287. 1990.
Pinus massoniana Lamb. var. shaxianensis Ecology
D. X. Zhou, Bull. Bot. Res. North-East. Forest. Inst.
Said to be growing on hills, without further details.
11 (3): 41. 1991.
Conservation
Description
This variety is poorly known but appears to occur in
Bark grey-brown towards base of trunk. Seed cones a limited area in the western part of Hainan Island,
ovoid or conical-ovoid when opened, 3–9 × 2.5–4 where it is probably declining due to deforestation.
cm. IUCN: EN (B1b)
 Pinus maximartinezii Rzed., Ciencia (Mexico) 23: at mid-cone, usually straight, transversely keeled,
17, t. 2. 1964. Type: Mexico: Zacatecas, Juchipila, with angular upper margin, those on the proximal
Pueblo Viejo, [“Cerro de Pinones”], J. Rzedowski and distal scales narrowly conical, often curved, dull
18258 (holotype MEXU). Fig. 229, 230 light brown or reddish brown, often resinous. Umbo
dorsal, obtuse-triangular or rhombic-pyramidal,
sometimes with a minute prickle, concolorous or
Etymology
grey-brown. Seeds oblong or ovoid-oblong, slightly
The species epithet commemorates the Mexican flattened, 20–28 × (8–)10–12 × 7–10 mm, wingless
botanist Maximino Martínez (1888–1964), who when the seed is free; integument ca. 2 mm thick,
studied the conifers of Mexico, but did not know very hard. Seedlings large, with 18–24 cotyledons;
708 this pine; a concomitant meaning refers to the size juvenile leaves curved, flattened, ca. 8 cm long,
(Latin: maxime) of the cones and seeds. ­silvery-blue, persisting well beyond the develop-
ment of adult leaves, sometimes up to 20 years.
Vernacular names
Distribution
piñon real (Mexico)
Mexico: S Zacatecas (Sierra de Morones).
TDWG codes: 79 MXE-ZA
Description
Trees to 5–10(–15) m tall, d.b.h. to 40–50 cm; trunk
Ecology
monopodial, short, branching low, often contorted
or curved. Bark relatively thick on lower trunk, thin The rock of the mountain where P. maximartinezii
elsewhere, smooth but eventually rough, tessellated occurs is in part sandstone or limestone and also
into square, ca. 10 cm wide plates; outer bark grey. metamorphic, the soils are very rocky and shallow.
Branches long, ascending to erect in the top, but Precipitation is probably ca. 700–800 mm annually,
lower branches spreading, those bearing cones be- virtually restricted to four months in the summer.
coming pendulous. Shoots glabrous or with puberu- Pinus maximartinezii is virtually the only pine here,
lent bases of fascicles, initially glaucous or greyish but a few scattered individuals of P. leiophylla var.
green, turning orange-brown to grey. Cataphylls ca. chihuahuana have been seen. Its altitudinal range
5 mm long, narrowly triangular-caudate, recurved. is 1800–2400 m a.s.l. Abundant are various large
Vegetative buds small, ovoid-conical; terminal buds leaved species of deciduous Quercus, e.g. Q. macro­
5–8 mm long, resinous. Fascicle sheaths 7–8 mm phylla, which are bare during the long dry season
long, light brown; the outer scales deciduous, the from September to May. Fires occur regularly in
inner scales recoiling, semi-persistent, forming a the region in all vegetation types; it is not known
small rosette at base of fascicle. Leaves in fascicles whether this species is adapted well to reseed itself
of 5, very rarely 3 or 4, persisting two years, straight, after fire. Pollen dispersal is usually in May-June;
lax, 7–11(–13) cm long, 0.5–0.7 mm wide, with entire the ovuliferous cones take 18–24 months to reach
margins, acute, glaucous green, in some trees green, maturity, and perhaps longer for the seeds to ripen
the two adaxial faces often whitish. Stomata only on fully, which mostly remain in the cones. Squirrels
adaxial faces. Pollen forming an elongated ‘spike’, are capable of biting off the apohyses to reach the
ovoid-oblong, 8–10 mm long, yellowish. Seed cones seeds and probably store them. They, and probably
lateral, solitary on ultimate branches, on a very short also birds, may play a crucial role in effective seed
peduncle or almost sessile. Mature cones broadly dispersal, but this has not been investigated to date.
ovoid-truncate, (15–)17–25(–27) × 10–15 cm when
the scales have parted. Seed scales parting slowly,
Conservation
usually insufficiently wide to release the seeds, very
thick woody, rigid, obtrullate, up to 50 mm wide The species is considered Endangered due to fire and
below the apophysis, of similar shape around the grazing hazards resulting in few seedlings that suc-
cone but differentiating from base to apex, with ceed in establishing themselves. Intensive harvest-
deep seed cavities on adaxial side. Apophyses very ing of cones and seeds may also diminish its chances
prominent, 35–50 × 20–35 mm, rhombic-pyramidal of reproduction. Although there is an awareness
amongst botanists and foresters in Mexico of its im- part of trunk, with relatively smooth, longitudinal
portance, current in situ protection (fire prevention plates and deep longitudinal fissures, grey-brown.
and fighting) seems inadequate to ensure long term Branches spreading or ascending, forming a py-
preservation of this interesting narrow endemic. ramidal crown in young trees and an open or dense,
Seed collections have been made in recent years to rounded crown in mature trees. Shoots uni-nodal,
ensure ex situ conservation programmes. The land green or light brown, rarely glaucous. Cataphylls
on which these pines grow is privately owned by vil- subulate-caudate, soon reflexed, 10–15 mm long.
lagers, who have an interest in the seed harvests as Vegetative buds ovoid-conical; terminal bud 15–20
well as cattle grazing. mm long; lateral buds smaller, not resinous. Fascicle
IUCN: EN (B1c) sheaths 15–25(–30) mm long, persistent, (lustrous)
grey-brown to grey. Leaves in fascicles of 5, rarely 709
4 or 6, persisting 2–2.5 years, slender, lax, drooping,
Uses
sometimes pendulous, 20–35 cm long, 0.6–1.0(–1.1)
Like other ‘piñon’ (pinyon pines) in Mexico, this mm wide, with serrulate margins, acute, (yellowish)
species is of local importance for its edible seeds, light green to glaucous green. Stomata on all faces
which are harvested by local people and marketed of leaves. Pollen cones densely clustered, cylindri-
in the region. Due to its low stature and branching cal, 3–4 cm × 5–8 mm when full grown, light pink-
of the trunk, its timber is not used. In Mexico, it is ish brown, turning darker. Seed cones subterminal,
sometimes planted as an ornamental tree; elsewhere solitary or in pairs on distinct, stout, curved pedun-
it is only grown in a few botanic gardens (e.g. at the cles which fall with cones. Mature cones narrowly
University of California in Berkeley) and research ovoid to ovoid-attenuate when closed, more or less
nurseries; its horticultural merits could be greater ovoid, slightly curved, with an obliquely flattened
than that since it is not too difficult to grow from base when opened, (4–)5–10(–12) × (3–)4–8 cm
seed in the nursery. Young trees retain an attractive when open. Seed scales thin woody, flexible, oblong,
blue juvenile foliage for several years. straight, spreading wide or often reflexed. Apophy-
sis flattened or slightly raised and then transversely
keeled, 8–15 mm wide, irregularly rhombic to pen-
Pinus maximinoi H. E. Moore, Baileya 14: 8. 1966. tagonal in outline, variably light brown. Umbo dor-
Pinus tenuifolia Benth., Pl. Hartweg.: 92. 1842, non sal, raised, curved, transverse-rhombic in outline.
Salisb. (1796); Pinus douglasiana Martínez var. Seeds obliquely ovoid, slightly flattened, 4–6 × 3–4
maximinoi (H. E. Moore) Silba, Phytologia 68: 50. mm, ochraceous or dark brown, with or without
1990. Type: Guatemala: [“In montibus para ruptis dark spots. Seed wings oblong, with a straight side,
que Canales dictis, nec non ad pagum Chinanta widest near the middle, 13–22 × 4–8 mm, yellowish
prope Guatemala, et in summo jugo Choacas prope brown, translucent.
Salama.”], C. T. Hartweg 620 (holotype K).
Distribution
Etymology
Mexico: Sinaloa, Jalisco, Michoacán, México,
This species was named after the Mexican botanist Hidalgo, Tlaxcala, Puebla, Veracruz, Guerrero, Oax-
Maximino Martínez (1888–1964), who studied the aca and Chiapas; Guatemala; Honduras; El Salvador
pines of Mexico. and NW Nicaragua.
TDWG codes: 79 MXC-ME MXC-PU MXC-TL MXE-
HI MXG-VC MXN-SI MXS-CL MXS-GR MXS-JA MXS-
Vernacular names
MI MXS-NA MXS-OA MXT-CI 80 ELS GUA HON NIC
Thin-leaf pine; ocote, pino canís (Mexico, Hondu-
ras)
Ecology
Pinus maximinoi is a species with a wide ecologi-
Description
cal amplitude, occurring from wet subtropical for-
Trees to 20–40(–50) m tall, d.b.h. to 70–90(–100+) est, where it is a gap pioneer, well up into the cooler
cm; trunk monopodial, straight. Bark thick on lower cloud forests on high mountains in Mesoamerica. In
 Mexico it also occupies drier sites as a constituent of Pinus merkusii Jungh. & de Vriese, in De Vriese,
pine or oak-pine forest or woodland. Its altitudinal Pl. Nov. Ind. Bat.: 5, t. 2. 1845. Type: Indonesia:
range is great: (450–)600–2800 m, with an optimum Sumatera, Aceh, [“hab. in provincia Battarum in
at 900–1800 m a.s.l. In the NW of its range the spe- montibus et rupibus Tanna Hurung et Tobak”],
cies occurs between 1500–2800 m. It occurs on a va- F. W. Junghuhn & W. H. de Vriese 1 (holotype L).
riety of soils under various climatic conditions; in
Mesoamerica annual precipitation ranges from ca. Pinus merkusii Jungh. & de Vriese subsp. ustulata
900–2500 mm, with the wettest conditions on the Businský, Acta Pruhon. 88: 7. 2008.
Atlantic and Pacific slopes of the mountains. Un-
der these conditions it occurs frequently with Pinus
Etymology
710 ­tecunumanii and Liquidambar styraciflua, at lower
altitudes with P. oocarpa. In secondary broad-leaved This species was named after the Lieutenant Gov-
forest other pines may join: P. devoniana, P. pseudo­ ernor-General of the Dutch East Indies Hendrik
strobus, and pine woodland may prevail under a re- Merkus de Kock, who held this post in 1826–1830.
gime of grazing or burning, with the undergrowth
dominated by grasses or the fern Pteridium aqui­
Vernacular names
linum. In Central Mexico, it grows at the higher
sites with Abies religiosa, P. ayacahuite, P. patula, Merkus’s pine, Mindoro pine; Tapulaw (Philippines)
P. pseudostrobus, P. douglasiana, and often Quercus
in mixed pine or oak-pine forest, where precipita-
Description
tion is more moderate but the seasonal temperature
range greater, with some frosts occurring in winter. Trees to 45(–70?) m tall; trunk to 2 m d.b.h., erect,
straight. Bark thin and almost smooth, reddish brown
or dark brown, or thick and scaly, rough, breaking
Conservation
into numerous small, dark grey plates. Branches
IUCN: LC spreading and ascending, forming an open, broadly
conical or irregular crown. Foliage branches stout,
glabrous, brown or dark brown. Buds cylindrical;
Uses
terminal bud 1.5–2 cm long, all without resin; cata-
Thin-leaf pine is an important timber tree in most phylls brown. Leaves in fascicles of 2, held by a per-
of its range, where it is exploited with other tall sistent, 12–18 mm long, reddish brown basal sheath,
growing pines. It has also been the subject of experi- falling in their second year, long and very slender,
mental forestry in various subtropical and tropical pliant, 15–20 cm long, straight or slightly contorted,
countries under programmes such as those initiated 1 mm wide, dull green; margins minutely serrulate;
by the Central America & Mexico Coniferous Re- apex short acute; stomata in numerous lines on all
sources Cooperative (CAMCORE). The wood is rel- surfaces. Pollen cones clustered, spirally arranged,
atively soft and light and easily worked and is used erect, cylindrical, 1.5–2.5 cm long, 5 mm wide. Seed
for construction (beams and planks), carpentry and cones sub-lateral on new shoots, usually solitary,
joinery, crates, containers and boxes, woodware, sometimes in pairs, on ca. 1 cm long, stout pedun-
tool handles, and matches, as well as various types of cles, spreading at nearly right angles from branch,
board, plywood and pulp. Thin-leaf pine is scarcely oblong-conical when closed, (5–)6–10(–11) cm long,
known in cultivation as an ornamental tree, but it widening to 4–8 cm and becoming broadly ovoid
has been imported as a forestry tree in Nepal as well with spreading scales. Seed scales woody, rigid, ob-
as in southern Africa and in Colombia, mostly in the long, twice as long as wide, ca. 3 × 1.2–1.5 cm at mid
context of experimental tree breeding programmes cone in the larger cones, light reddish brown; apo-
with the objective of producing fast growing planta- physes rhombic or unequally pentagonal in outline,
tion trees for industry. These trials have as yet not raised and strongly transversely keeled, radially stri-
led to large scale economic use. ated or ridged, lustrous red-brown to dark brown;
umbo dorsal, more or less central, slightly sunken
or flat to obtuse, unarmed. Seeds obovoid, 5–7 mm
long, 4.5 mm wide, slightly appressed, grey-brown, resin. Young planted trees are better for tapping than
with a 20–25 mm long persistent wing. ‘old growth’ trees in natural stands. Indonesia is a
major producer of turpentines distilled from this
resin. In the Philippines, this species is tapped to-
Taxonomic notes
gether with P. kesiya, which is indigenous on these
In most of the previous handbooks or manuals on islands, but not in Indonesia. When trees have
conifers, Pinus merkusii has been treated in the grown beyond good yield of resin, their wood is har-
broad sense to include populations occurring in vested for the pulp industry to manufacture paper,
mainland SE Asia, here considered to belong to a process which allows final extraction of the resin
P. latteri Mason. See for a discussion on this issue in the wood. If well managed, these plantations are
and the (purported) differences between the two a renewable resource and can assist in the preserva- 711
taxa under P. latteri. tion of the natural stands of P. merkusii. The wood
of higher grade is also used in house construction,
panelling and furniture making. This species seems
Distribution
not to have been planted as an ornamental, but it is
Malesia: N Sumatera, from around Lake Toba present in a few botanic gardens in Indonesia and
NE along mountains; Philippines, in Luzon and the Philippines.
­Mindoro.
TDWG codes: 42 PHI SUM
Pinus monophylla Torr. & Frém., in Frémont, Rep.
Exped. Rocky Mts.: 319, t. 4. 1845, [“monophyllus”].
Ecology
Pinus cembroides Zucc. subsp. monophylla (Torr. &
Pinus merkusii occurs in mountainous regions and Frém.) E. Murray, Kalmia 12: 22. 1982. Type: USA:
forms more or less open pine woods or pine savan- California, [“collected in northern California”],
nas influenced by periodic grass fires. This eco­ J. C. Frémont 367 (holotype NY). Fig. 231, 232
system is much influenced by man and may even
have been created by people over thousands of Pinus edulis Engelm. var. fallax Little, Phytologia 17:
years of occupation. Like its vicariant species on the 331. 1968; Pinus californiarum D. K. Bailey subsp. fal­
SE Asian mainland, P. latteri, a ‘grass stage’ is report- lax (Little) D. K. Bailey, Notes Roy. Bot. Gard. Edin-
ed to occur as an adaptation to these grass fires in burgh 44: 279. 1987; Pinus monophylla Torr. & Frém.
the seedling to sapling stages of its development. In var. fallax (Little) Silba, Phytologia 68: 54. 1990;
Sumatera, this is the only pine that crosses the equa- ­Pinus ×fallax (Little) Businský, Acta Pruhon. 88: 11.
tor into the southern hemisphere. 2008.
Pinus californiarum D. K. Bailey, Notes Roy. Bot.
Gard. Edinburgh 44: 278. 1987; Pinus monophylla
Conservation
Torr. & Frém. var. californiarum (D. K. Bailey) Silba,
Pinus merkusii is threatened by over-exploitation, Phytologia 68: 54. 1990.
habitat degradation and overgrazing. Around Lake
Toba in northern Sumatera historical decline has
Etymology
fragmented the population and exploitation has pro-
gressed to the NE from there. In the Philippines the The species epithet monophylla refers to the single
much smaller populations are now also fragmented. leaf (needle) in a fascicle.
IUCN: VU (B2c-e)
Vernacular names
Uses
Singleleaf Pinyon pine
Merkus’s pine has been extensively planted through-
out Indonesia (where it is only indigenous in north-
Description
ern Sumatera) by the Dutch in colonial times.
Indonesian foresters have continued this practice as Trees or a large shrubs to 15–20 m tall, d.b.h. to
it is the country’s most important producer of pine 40–50 cm. Trunk monopodial, usually short, low
 branched, straight or contorted. Bark thick, rough woodland in this area; the most common juniper
and scaly, with shallow, fissures, exfoliating in small is Juniperus osteosperma, in the NW replaced by
plates; outer bark reddish brown to grey. Branches J. occidentalis. At higher altitudes P. ponderosa can
spreading or ascending, forming a wide spread- be mixed in as an indicator of the transition to tall
ing, rounded but irregular, open crown often ex- pine forests; in the White Mountains of California
tending low above the ground. Shoots short, stout, P. monophylla has been found with P. longaeva at
orange-yellow, turning grey after 1–2 years. Cata- 3000 m. Its altitudinal range is from 950 m to 3000
phylls short, triangular, rigid and spreading. Vegeta- m a.s.l. The undergrowth is dominated by sagebrush
tive buds ovoid-conical, acute; terminal bud 10–15 (Seriphidium spp. [Artemisia]) and numerous other
× 5–7 mm; lateral buds smaller, (slightly) resinous. xerophytic shrubs are common. In southern Cali-
712 Fascicle sheaths of young fascicles to 11 mm long, fornia and Baja California, P. monophylla is locally
soon disintegrating in recoiling but almost simul- common in, or just above, a high chaparral zone but
taneously deciduous scales of which only the basal does not form extensive Pinyon-Juniper woodland.
parts persist (not forming rosettes). Leaves in fasci- Other common pines here are P. quadrifolia and
cles of 1, rarely 2, persisting 4–8 years, rigid, curved P. jeffreyi, the latter species extending to higher al-
at least near base, (2–)2.5–6 cm long, 1.2–2.2(–2.5) titude. Juniperus californica and Quercus turbinella
mm wide, in rare fascicles of 2 with entire margins, commonly occur with P. monophylla in areas with
­acuminate-pungent, usually lustrous, green, grey- granite rock outcrops and on steep slopes. Annual
green or glaucous green. Stomata around the leaves, precipitation is low to moderate from 200–600 mm,
with 12–25 lines in grooves. Pollen cones ovoid-glo- highly variable, in California concentrated in the
bose to short cylindrical, up to 10 mm long when winter and part of it as snow at higher altitudes. Se-
shedding pollen, initially purplish red, then yellow- vere frost can occur in the Great Basin, which has a
ish. Seed cones sub-terminal, solitary or in whorls of continental climate.
2–4 on slender, 5–10 mm long peduncles which are
deciduous with cones. Mature cones ovoid-globose
Conservation
to globose when closed, irregular when opened, of-
ten somewhat wider than long, 4–6 × 4.5–7 cm when IUCN: LC
open. Seed scales parting widely, spreading but
not reflexed, moveable, irregular, concavo-convex,
Uses
with 1–2 deep seed. Apophysis thick woody, raised,
­pyramidal or obtuse conical, transversely keeled, re- Due to its irregular shape and slow growth, this tree
curved or straight, lustrous, ochraceous to yellowish is not used for lumber; locally it is used as firewood.
brown, often resinous. Umbo dorsal, flat or obtuse- The edible seeds are locally harvested and sold on
pyramidal, rhombic in outline, centrally indented. village markets or alongside the highways. In coun-
Seeds obliquely obovoid, 13–18 × 8–12 mm, integu- tries with hot summers this would be an excellent
ment thin (0.3–0.5 mm), greyish brown to grey. small pine for amenity planting, but it is little used
Megagametophyte (‘endosperm’) white. Seed wings and only present in a few collections. In the USA,
absent from the free seed. where pines are grown for Christmas trees, it can
be put to this use, but it is slow growing and needs
much clipping to attain the desirable shape.
Distribution
USA: Arizona, California, Idaho, Nevada, Utah;
NW Mexico: Baja California Norte. Pinus montezumae Lamb., Descr. Pinus, ed. 8°, 1:
TDWG codes: 73 COL IDA 76 ARI CAL NEV UTA 79 39, t. 22. 1832.
MXN-BC
Etymology
Ecology
Montezuma II was the Aztec emperor killed in a riot
This species of pinyon pine is common on the dry in Tenochtitlán (Mexico City) in 1520 shortly before
mountain slopes of the Great Basin. It is the ma- the Spanish conquest.
jor component of the extensive Pinyon-Juniper
 Querétaro, Hidalgo, Morelos, Tlaxcala, Puebla, Cen-
Vernacular names
tral Veracruz, Guerrero, Oaxaca and Chiapas; in
Montezuma pine, Rough-branched Mexican pine; Guatemala in the highland departments.
ocote blanco, pino de montezuma, pino real (Mex- TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
ico). PU MXC-TL MXE-NL MXE-QU MXE-HI MXE-TA
MXE-ZA MXG-VC MXS-CL MXS-GR MXS-JA MXS-MI
MXS-NA MXS-OA MXT-CI 80 GUA
Description
Trees to 20–30 m tall, d.b.h. to 1 m; trunk mono­
Ecology
podial, erect. Bark thick, scaly, breaking into
numerous, relatively small, irregular, slightly elon- Pinus montezumae occurs in a wide range of edaph- 713
gated plates divided by shallow fissures, dark brown, ic and climatic conditions throughout the moun-
weathering grey-black. Branches spreading wide, tainous regions of Central and S Mexico and the
assurgent towards ends, forming a dense, usually Guatemalan highlands. In Nuevo León it reaches
broad, rounded crown. Shoots very rough and scaly, from semi-arid Pinyon-Juniper woodland up to
brown, sometimes glaucous. Cataphylls ca. 15 mm cold temperate mixed conifer forest. Its altitudinal
long, subulate, recurved or reflexed, scarious, dark range is also great, from (1200–)2000–3200(–3500)
brown, weathering grey. Vegetative buds ovoid; m overall, with occurrences below 2000 m mainly
terminal bud 15–30 mm long; lateral buds smaller, in Nayarit, Nuevo León and Tamaulipas. It is most
not resinous. Fascicle sheaths (20–)25–35 mm long, abundant and best developed in the temperate zone
1.5–2.5 mm wide, remaining tight, weathering grey at ca. 2400–2800 m a.s.l., with annual precipitation
to black, sometimes resinous. Leaves in fascicles of exceeding 800 mm, most of it falling from June
(4–)5 (rarely 3 or 6), persisting 2–3 years, slender, through September. Throughout its wide geographi-
flexible and drooping, or lax to rigid and spreading, cal range, it occurs together with many other tree
straight, (15–)20–35(–40) cm long, 0.8–1.3 mm wide, species, mainly in pine-oak and mixed pine for-
with serrulate margins, acute or pungent, (light) ests, but also with Abies religiosa, A. guatemalen­
green to glaucous green. Stomata on all faces of sis and Cupressus lusitanica. Many of these forests
leaves. Pollen cones cylindrical, 20–40 × 6–8 mm, have been depleted or even turned into small scale
initially pink or purplish, turning reddish brown. farms with groups of scattered trees, among which
Seed cones subterminal, solitary or in whorls of 3–6, P. ­montezumae is often present.
spreading or recurved on short, stout peduncles.
Mature cones variable, from ovoid or symmetrical
Uses
ovoid-oblong to ovoid-attenuate, often curved, with
a more or less flattened base when opened, 8–20 × Pinus montezumae is exploited as a timber tree
5–10 cm when open. Seed scales parting to release throughout its range. The timber is heavy and
the seeds, the proximal, infertile scales remaining on strong and in demand for constructional purposes,
branch when cone falls, thin or thick woody, oblong, plywood, parquet flooring, and furniture. Together
straight or curved. Apophysis flat or raised especial- with the wood of other pines it is pulped for the
ly on proximal scales, if raised with a transverse keel, paper industry. Locally, especially in the southern
(transverse-)rhombic in outline, sometimes irregu- part of its range, this (and other) species, growing
lar, often rugose or radially striate, in various shades often near habitations among fields as remnants of
of ochraceous to brown. Umbo dorsal, depressed, more contiguous forests, has its branches cut off due
flat or raised, obtuse or terminating in a small to the incessant demand of firewood of the rural
prickle. Seeds obliquely ovoid, 5–7 × 4–5 mm, light population. Since pines rarely regenerate (only one
brown, often with dark spots. Seed wings oblong, Mexican species is known to do so) and none can
18–28 × 7–12 mm, light brown with dark stripes. be coppiced or pollarded indefinitely, this use will
be detrimental and lead to the demise of the trees.
­Pinus montezumae is a very handsome pine grown
Distribution
in horticulture for large gardens and parks; it is
Mexico: Nuevo León, SW Tamaulipas, Nayarit, S Za- moderately hardy depending on provenance.
catecas, Jalisco, Michoacán, México, Distrito ­Federal,
 2 varieties are recognized:
Conservation
Pinus montezumae Lamb. var. montezumae. Type: IUCN: LC
Mexico: Veracruz, [“inter Cruz Blanca et Jalacingo,
reg. frig. Nov. 28”], C. J. W. Schiede & F. Deppe s.n.
(lectotype BM). Pinus monticola Douglas ex D. Don, in Lambert,
Descr. Pinus, ed. 8°, 2: p. s.n. inter 144 et 145. 1832.
Pinus montezumae Lamb. var. mezambrana Carva- Pinus strobus L. var. monticola (Douglas ex D. Don)
jal, Phytologia 59: 138. 1986 [“mezambranus”]. Nuttall, N. Amer. Sylva 2: 118. 1849; Pinus strobus
L. subsp. monticola (Douglas ex D. Don) E. Murray,
714 Kalmia 12: 23. 1982. Type: USA: Washington, [“high
Description
mountains at the Grand Rapids”], D. Douglas s.n.
Leaves in fascicles of (4–)5(rarely 3 or 6), variable, (holotype not located, isotype K).
slender and drooping or more rigid, 1.0–1.3 mm
wide. Seed cones variable in shape and size, broadly
Etymology
ovoid, ovoid-oblong or ovoid-attenuate, 8–20 × 5–10
cm when open. Apophysis raised, especially towards The species epithet monticola means ‘inhabitant of
base of cone, transversely keeled, often rugose; mountains’.
umbo raised, sometimes with a small prickle.
Vernacular names
Distribution
Western white pine
Central Mexico south to Chiapas.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
Description
PU MXC-TL MXE-HI MXE-NL MXE-QU MXE-TA
MXE-ZA MXG-VC MXS-GR MXS-JA MXS-MI MXS- Trees to 60–70 m tall; trunk to 2.5–3.3 m d.b.h.,
NA MXS-OA MXT-CI growing to a straight and columnar bole, scarcely
tapering in lower half of its length. Bark smooth in
young trees, on large trunks becoming scaly, exfolia-
Conservation
ting in small or large, rectangular plates, cinnamon
IUCN: LC to grey-brown. Branches numerous, relatively short,
spreading or down-curved, forming a conical but
Pinus montezumae Lamb. var. gordoniana (Hartw. in old trees rounded crown. Foliage branches slen-
ex Gordon) Silba, Phytologia 68: 55. 1990. Pinus der, smooth, young shoots puberulent with brown,
gordoniana Hartw. ex Gordon, J. Hort. Soc. London downy hairs, later glabrous, grey-green or reddish
2: 79. 1847. Type: Illustration in J. Hort. Soc. 2: fig. tan, becoming grey. Buds small, with the terminal
on p. 80. 1847 (lectotype). bud to 6 mm long, ovoid-ellipsoid, orange-brown
or red-brown, slightly resinous. Leaves in fascicles
of 5, persisting 3–4 years, held in a soon deciduous
Description
sheath of light orange-brown flimsy scales, spread-
Leaves in fascicles of (4–)5, slender and droop- ing, (4–)5–10 cm long, straight, slender, flexible,
ing, 0.8–1.0 mm wide. Seed cones ovoid-oblong or 0.7–1 mm wide, slightly twisted, glaucous green
ovoid-attenuate, 10–15 × 5–7 cm when open. Apoph- with lines of stomata on the two adaxial faces only;
ysis flat, especially towards base of cone, with a weak margins minutely serrulate; apex acute. Pollen cones
transverse line, often smooth; umbo flat or obtuse. ellipsoid to short cylindrical, 10–15 mm long, yellow.
Seed cones clustered, erect at first, becoming pen-
dulous as they grow on 20 mm long peduncles, soon
Distribution
falling after seed dispersal, 10–25(–30) cm long,
Central and S Mexico to Oaxaca. variable in length on the same tree, oblong cylindri-
TDWG codes: 79 MXC-PU MXE-QU MXG-VC MXS- cal becoming ellipsoid cylindrical when open, sym-
GR MXS-JA MXS-MI MXS-NA MXS-OA metrical or nearly so. Seed scales thin woody, more
or less ­flexible, small scales near the cone base re- occurs on a diversity of soil types. Pinus monticola
flexed, larger scales more or less straight or with an can form extensive pure stands, but old growth for-
upturned apophysis, extremely resinous. Apohysis ests are usally mixed conifer forests with several oth-
more or less rhombic in outline on central scales, er species. Depending on region and site conditions,
tan coloured (lower part of scales purplish brown), the most common are Pseudotsuga menziesii, Pinus
narrowing to an obtuse terminal umbo. Seeds contorta, P. ponderosa, P. lambertiana, P. jeffreyi,
broadly obovoid to deltoid, flattened, 5–8 mm long, Tsuga heterophylla, Larix occidentalis, Abies grandis,
red-brown; wing 20–25 mm long. A. lasiocarpa, A. magnifica, A. concolor, A. procera,
Thuja plicata, Calocedrus decurrens, Picea sitchen­
sis, P. engelmannii, and Tsuga mertensiana. More
Taxonomic notes
local are Chamaecyparis lawsoniana, Pinus balfou­ 715
The morphological differences between Pinus mon­ riana, P. flexilis, and P. albicaulis. Taxus brevifolia is
ticola and P. strobus are small, but a few are dis- an understorey conifer. Broad-leaved trees are less
continuous. The bark of large boles in P. monticola common and usually restricted to sites with better
breaks into plates, but does not become longitudi- moisture and nutrient availability. These are some of
nally furrowed as in P. strobus. The basal scales of the the most diverse conifer forests in the world. Pinus
seed cones of P. monticola are recurved, in P. strobus monticola is an early seral species in this assembly
they are incurved or straight. There are some minor and requires disturbance of the forest (by fire, storm
differences in leaf apices and in the colour of the or felling) to regenerate.
ripe and dry scales of the seed cones, but they seem
less consistent. It could well be argued (see also the
Conservation
case of P. strobus var. chiapensis), that here we have
an eastern (P. strobus) and a western (P. monticola) IUCN: LC
vicariant population of a single species, which has
evolved a few minor distinctions recognizable at
Uses
the rank of subspecies. However, some phylogenetic
analyses based on DNA sequence data indicated a Western white pine is an important timber tree
closer relationship between P. monticola and P. lam­ which yields high quality, straight-grained wood of
bertiana than between P. monticola and P. strobus. good dimensions and strength. It is used for interior
These results support the separation of P. monticola construction and paneling, doors and windows of
and P. strobus as distinct species. houses and plywood; other applications of the wood
are furniture, matches and tooth picks. Matches have
become a major application of the wood of smaller
Distribution
trees of this species produced in secondary growth
W North America, from British Columbia to Cali- forests. Its wood does not produce sugary exudates
fornia, most widespread in the north of its range. (resin containing a high content of saccharine) as
TDWG codes: 71 ABT BRC 73 IDA MNT ORE WAS in P. lambertiana, a closely related species of the
76 CAL NEV American West. Western white pine grows rapidly
and can be easily regenerated after felling and grows
straight up forming tall stems. As with P. lamber­
Ecology
tiana, its susceptibility to Western white pine blister
Pinus monticola has its greatest extent in the cool rust (Cronartium ribicola, Basidiomycota) makes it
maritime Pacific Northwest of the USA and Can- unsuitable as an exotic timber tree in Europe and
ada, its southward spread is limited to the western other parts of the temperate climate zones where
slopes of the high mountain ranges. In interior Brit- this pathogen occurs. It is present in a few arboreta
ish ­Columbia it is restricted to the valleys and basins and landscape parks, but uncommonly used in hor-
and does not ascend beyond 450 m a.s.l. while in ticulture and only a few cultivars are known from
the Sierra Nevada of California it climbs to beyond this species.
3000 m. In the interior and in the south, this species
 Pinus morrisonicola Hayata, Gard. Chron., ser. 3, ing more ovoid. Seed scales thin woody, somewhat
43: 194. 1908. Pinus parviflora Siebold & Zucc. var. flexible, with basal smaller scales mostly recurved
morrisonicola (Hayata) C. L. Wu, Acta Phytotax. and larger scales nearly straight, 3–3.5 cm long,
Sin. 5 (3): 141. 1956. Type: Taiwan: [“Shokakulin”], 1.5–2 cm wide, cuneate from their base; apophyses
C. Owatari s.n., 21 Jan 1898 (holotype TI). rhombic, middle portion thickened, longitudinally
grooved, becoming lustrous brown; umbo terminal,
Pinus formosana Hayata, J. Linn. Soc., Bot. 38: 297. obtuse, slightly upturned. Seeds ellipsoid-ovoid or
1908. narrowly ovoid, 7–10 mm long, 5–6 mm wide; wing
Pinus uyematsui Hayata, Icon. Pl. Formos. 3: 192, t. 15–20 mm long, 5–8 mm wide, light brown.
35. 1913.
716 Pinus ×hayatana Businský, Willdenowia 34 (1): 245. Distribution
2004. [putative hybrid between P. uyematsui and
Taiwan.
P. morrisonicola]
TDWG codes: 38 TAI

Etymology Ecology
The species epithet means ‘growing on Mt. Morri- Pinus morrisonicola occurs in the mountains of Tai-
son’, the highest mountain in Taiwan, now known wan on rocky ridges and spurs at various altitudes
as Yu-Shan. between ca. 300 m and 2300 m a.s.l. Occasionally
it occupies slopes where the forest canopy has been
opened, e.g. due to rock slides, so it can compete for
Vernacular names
a time with broad-leaved trees until the forest suc-
Taiwan white pine; tai wan wu zhen song (Chinese) ceeds to a closed canopy again.

Conservation
Description
IUCN: LC
Trees to 30–35 m tall; trunk to 1.5 m d.b.h., usually a
straight, columnar bole. Bark smooth in young trees
Uses
and in upper part of crown, becoming rough and
scaly on trunk, flaking in thin plates, grey to dark Taiwan white pine is a minor timber tree for lo-
grey. Branches whorled in young trees, later spread- cal use, with wood properties similar to those of
ing and ascending, stem sometimes forked near the P. parviflora and other E Asian white pines. It is bet-
top, forming a conical to rounded crown, on exposed ter known in Asian horticulture, especially in the
sites often flattened and open. Foliage branches slen- art of bonsai culture, and particularly in China. In
der, new shoots initially yellowish pubescent, soon Western horticulture it is uncommon and prob-
glabrous or with remnant hairs in grooves, reddish ably often confused with the much more commonly
brown. Buds small, ovoid, not resinous, pale brown. used Japanese white pine (P. parviflora), to which it
Leaves in fascicles of 5, held by deciduous sheaths is closely related, but differs in the longer leaves and
with soon falling thin scales, persisting 3–4 years, larger seed cones. As an endemic species of Taiwan,
sometimes longer, curved and slightly twisted, 4–9 it should not be difficult to establish its identity in
cm long, slender and flexible, 0.6–1 mm wide, trian- cultivation, provided that proper records of prov-
gular in cross-section, green; stomata in prominent enance were kept with the plants in tree nurseries.
white lines on the two adaxial surfaces; margins
minutely serrulate; apex acute. Pollen cones in small
clusters near base of new shoots, spirally arranged, Pinus mugo Turra, Giorn. Italia Sci. Nat. 1: 152.
ovoid-oblong to cylindrical, 1.5–2.5 cm long, yellow 1764.
turning light brown. Seed cones in whorls of 3–4 on
short, stout, curved peduncles, erect at first, becom-
Etymology
ing pendulous when growing, (narrowly) ovoid-
ellipsoid, 6–11 cm long, green or glaucous, very The species epithet uses the Italian vernacular name
resinous, with opened scales to 5–6 cm wide, becom- for this pine.
 may still elude us as some recent papers seem to
Vernacular names
attest. A full synonymy following what was then in
Dwarf mountain pine, Mountain pine, Mugo pine; my view the most reasonable taxonomy by 2000 has
Bergkiefer (German); pin de montagne (French); been given in my World Checklist & Bibliography of
mugo (Italian); kosodrzewina (Polish). Conifers (Farjon, 1998, [2001]), several new names
had to be added since and it does not look as if it will
end there yet. The only subspecies recognized here is
Description
subsp. rotundata, which is usually a small erect tree.
Shrubs to 5 m tall, main branches ascending to The distinct asymmetric cones with prominent apo-
erect, sometimes a small erect tree. Bark breaking physes on the exposed side have been found in some
into small, rectangular plates and sometimes curling populations of shrubby forms and are not restricted 717
scales on larger stems, grey. Foliage branches slen- to the tree forms of the western Alps and Pyrenees.
der, smooth with prominent ribs, glabrous, at first The seed cones of P. mugo appear to be highly vari-
light greenish brown, later reddish brown to grey- able and too much taxonomic significance has been
ish black. Buds cylindrical, acute, strongly resinous, given to these variations.
10–15 mm long. Leaves in fascicles of 2, rarely 3, held
in 10–18 mm long, later much shorter, basal fascicle
Distribution
sheaths, densely set on branches, directed forward,
persisting (2–)4–9(–10) years, (2.3–)3–7(–8) cm Europe, in mountains from W Spain to Bulgaria and
long, slightly curved and rigid, often twisted, 1.5–2.2 Romania.
mm wide, dark or light green; margins minutely TDWG codes: 11 AUT-AU CZE-CZ CZE-SK GER POL
serrulate; apex more or less pungent; stomata in SWI 12 FRA-FR SPA-AN SPA-SP 13 ALB BUL ITA-IT
fine lines on all sides. Pollen cones clusterd at base ROM YUG-BH YUG-CR YUG-SE 14 UKR-UK
of new shoots, spirally arranged, short cylindrical,
10–12 mm long, 5 mm wide, yellow or red. Seed
Ecology
cones solitary or in whorls of 2–3 on short pedun-
cles below shoot apex, or sometimes more in a clus- There are two principal growth forms of Pinus mugo,
ter near its base and subsessile, patent or more or commonly recognised as distinct taxa: a shrub-like,
less reflexed, falling at maturity or persisting up to sometimes nearly decumbent form (subsp. mugo)
4 years, (obliquely) ovoid, 2–6 cm long, 2–4.5 cm and an erect tree (subsp. rotundata), which occupy
wide when opened, with an asymmetrical or sym- different habitats. The shrubby form grows on moun-
metrical, flattened base. Seed scales thin woody, tain slopes and ridges from about 1000 m to 2300
rigid, oblong, spreading wide, reddish or blackish m a.s.l. in the mountain ranges of Europe most ex-
brown; apophyses nearly flat to prominently raised, posed to storms associated with depression systems
gibbous or transversely keeled, on the outer, sun- in the North Atlantic. Especially in the Carpathi-
exposed side of cone sometimes larger and recurved ans, it forms dense mat-like thickets above montane
or longitudinally ridged, lustrous yellowish or red- forests dominated by Fagus or Picea; in the western
dish brown; umbo dorsal, small, depressed, flat or Alps and the Pyrenees the upright form (subspecies)
conical, central or excentric, armed with a minute, dominates on nutrient poor slopes. Pinus mugo in
often breaking prickle. Seeds small, 3–4 mm long; the eastern Alps may have replaced original Larch-
wing articulate, 10–13 mm long. Arolla pine woods which were disturbed by human
activities and grazing of their animals. The species
often occurs on dolomite limestone, but is in fact in-
Taxonomic notes
different to soil type; this prevalence probably has
Pinus mugo is probably the species of pine with the historical reasons (Ellenberg, 1988). While upright
longest list of synonyms. Central and East European stands of P. mugo subsp. rotundata can be fairly
botanists in particular have described the observed rich plant communities, the associated species with
variation in this pine as distinct species, subspecies, the decumbent subsp. mugo are much fewer due to
or varieties in almost every conceivable nomenclatu- harsh environmental conditions, such as exposure
ral combination. Despite several efforts at distentan- and long-lasting snow cover.
gling the taxonomy and nomenclature, a consensus
 Pinus mugo Turra subsp. rotundata (Link) Janch.
Uses
& H. Neumayer, Oesterr. Bot. Z. 91: 214. 1942. Pinus
The shrubby subspecies of Dwarf mountain pine has rotundata Link, Abh. Akad. Wiss. Berlin 1827: 168.
been used in some parts of northern Europe to stabi- 1830. Type: [locality not mentioned], 2/6 July 1981,
lize drifting sand dunes and as initial shelter belts for K. I. Christensen A4 (neotype B).
plantations with Scotch pine in similar sandy areas.
In horticulture it is mainly planted in spaces created Pinus uliginosa Neuman ex Wimm., Uebers. Arbe-
by roundabouts and other types of road intersec- iten Veränd. Schles. Ges. Vaterl. Cult. 1837: 96. 1838;
tion, both in Europe and in the USA. For gardens Pinus uncinata Ramond ex DC subsp. uliginosa
many cultivars that remain more dwarfish than the (Neumann ex Wimm.) Businský, Phyton (Horn,
718 subspecies mugo have been and are being selected, Austria) 46 (1): 132. 2006.
and some of these are suitable in larger rock gardens Pinus uncinata Ramond ex DC var. pseudopumilio
as they grow very slowly. For this reason this species Willk., Jahrb. Königl. Sächs. Akad. Forst- Landwirte
has also been used in bonsai culture. The tree form Tharandt 40: 218. 1861; Pinus mugo Turra subsp. ro­
(subsp. rotundata) is too uncommon and also grows tundata (Link) Janch. & H. Neumayer var. pseudo­
too slowly to be of importance as a timber tree. Its pumilio (Willk.) P. Schmidt, Sammelbl. Gebirgspfl.
horticultural interest is limited to arboreta, where (Karl-Marx-Stadt) 1982: 3. 1982.
it is often labeled as a distinct species (P. uncinata)
and can grow into an erect small tree. Hybrids have
Description
been described between subsp. mugo and subsp.
­rotundata, and such plants may also occasionally be Small, erect trees, rarely erect shrubs. Seed cones
in cultivation. asymmetrical; apophyses of outer (sun-exposed)
seed scales enlarged, curved, longitudinally ridged
2 subspecies are recognized: or keeled.

Pinus mugo Turra subsp. mugo. Types: Italy: Alto

Distribution
Adige, Vicenza, Monte Baldo, J. F. Séguier s.n. (lecto­
type [NIMES]) & J. B. Saint-Lager s.n. (epi­type G). Europe, Pyrenees (rare), Auvergne Mts., Alps, Erzge-
birge, Böhmerwald, Sudeten, NW Carpathians.
TDWG codes: 11 AUT-AU CZE-CZ CZE-SK GER POL
Description
SWI 12 FRA-FR 13 ITA-IT
Shrubs to 5 m tall, sometimes decumbent. Seed
cones more or less symmetrical; apophyses of seed Conservation
scales usually more or less equally developed around
IUCN: LC
cone, but variable.

Pinus muricata D. Don, Trans. Linn. Soc. London

Distribution
17: 441. 1836. Type: USA: California, San Luis
Europe: Pyrenees, Alps (rare in W Alps), Jura, Vosg- Obispo Co., Coon Creek, T. Coulter 712 (holotype
es, Erzgebirge, Böhmerwald, Sudeten, Carpathians, not located, isotype TCD). Fig. 233, 234
Rhodope Mts., Dinaric Alps, central Apennines.
TDWG codes: 11 AUT-AU CZE-CZ CZE-SK GER POL Pinus muricata D. Don var. borealis Axelrod ex
SWI 12 FRA-FR SPA-AN SPA-SP 13 ALB BUL ITA-IT Farjon, in Greuter (ed.), Names in Current Use 2,
ROM YUG-BH YUG-MA YUG-SE 14 UKR-UK Pinaceae: 136. 1993.
Pinus muricata D. Don var. stantonii Axelrod ex
Farjon, in Greuter (ed.), Names in Current Use 2,
Conservation
Pinaceae: 136. 1993.
IUCN: LC
Etymology
The species epithet (Latin adj. muricatim = ‘shaped
like a purple snail’) is emigmatic.
 Vernacular names Ecology
Bishop pine, Swamp pine Pinus muricata occurs from near sea-level to ca. 300
m a.s.l. in coastal areas. It grows within the chaparral
zone influenced by (summer and autumn) fog and
Description
winter rain, probably amounting to ca, 500 mm an-
Shrubs or small trees to 15(–25) m tall, d.b.h. to nually. Often brush fires sweep the area during the
20–50(–100) cm. Trunk monopodial, often branch- long, hot summers, killing stands of pines, but the
ing near the ground, erect or curved and crooked. serotinous cones are adapted to open quickly after
Bark rough and scaly, exfoliating, on larger trunks fire to release the seeds when the undergrowth has
with deep longitudinal fissures, dark brown to grey. been cleared away. Massive regeneration then quick- 719
Branches spreading or ascending, in shrubs often ly reoccupies these sites. This species grows in dry
assurging, forming an open, broad, irregular crown. sandy soils, on clay barrens and on swampy ground
Shoots multi-nodal, rough with large, short decur- or in peat bogs. The associated vegetation often con-
rent, persistent pulvini. Cataphylls ca. 10 mm long, sists of Adenostoma, Arctostaphylos, Seriphidium
subulate, curved or reflexed, scarious, brown. Veg- (Artemisia), Ceanothus, Heteromeles, Salvia, and
etative buds ovoid-acute; terminal bud 10–15 mm other shrubs.
long; lateral buds smaller, not or slightly resinous.
Fascicle sheaths 10–14 mm long initially, reduced to
Conservation
less than 10 mm on mature fascicles. Leaves in fas-
cicles of 2, persisting 2–3 years, straight or slightly IUCN: NT
curved, rigid, (7–)10–14(–16) cm long, 1.3–2.0 mm
wide, with serrulate margins, acute, light green or
Uses
dark green. Stomata on both faces of leaves. Pollen
cones oblong-cylindrical, 1.5–2 cm long, pink to red- Due to a limited range and small populations Bish-
dish. Seed cones sub-terminal, solitary or in whorls op pine is not a commercially valuable timber tree.
of 2–5 on stout, 5–10 mm long peduncles, reflexed, Its wood properties are of medium quality, with a
tenacious; mature cones seemingly sessile, opening coarse grain and much resin; its uses are therefore
only partly, serotinous, narrowly ovoid to ovoid- mainly heavy construction, beams, crates and some
attenuate when closed, or strongly asymmetrical, light construction, and carpentry or joinery, with
broadly ovoid, 5–7(–8) × 4–5(–6) cm when (half) the higher grades obtained from larger trees. This
open. Seed scales parting very slowly except those at species is well known in horticulture and amenity
base or lower half of cone, oblong, straight or slightly planting, it is often planted in parks and large gar-
curved, thick woody. Apophyses very variable, from dens, though not nearly as common as Pinus radiata,
slightly raised to extremely elongated, especially on which is also a ‘closed-cone’ pine from coastal and
upperside of cone, sometimes curved, up to 15 mm insular California. Bishop pine grows poorly on ca-
wide and 20 mm long, from dull, dark brown to lus- lacareous soils. Its botanical characters are quite var-
trous light brown. Umbo dorsal, if enlarged forming iable, yet no cultivars are known under this species.
a flattened, curved spine, armed with a sharp prickle
in obtuse forms, 2–10 mm long. Seeds obliquely
ovoid, 5–6 × 3–4.5 mm, grey to black. Seed wings Pinus ×neilreichiana Reichardt, Verh. Zool.
14–18 × 5–8 mm, yellowish brown to grey-brown. Bot. Ges. Wien 26: 461. 1876. Type: Austria:
Niederösterreich, Baden, Grossau, along footpath
to Pottenstein, [?] Reichardt s.n. (lectotype C).
Distribution
USA: California (Santa Cruz Island and Santa Rosa
Etymology
Island); Mexico: Baja California Norte.
TDWG codes: 76 CAL 79 MXN-BC The epithet commemorates the German botanist
August Neilreich (1803–1871).
 Vernacular names Description
None are recorded. Trees to 5–10 m tall, sometimes bushy, d.b.h. to
15–30 cm. Trunk monopodial, usually straight,
erect, some trees branching low near the ground.
Description
Bark thin, smooth, only on lower part of larger
This natural hybrid is morphologically closest to trees becoming scaly, ash-grey, with darker, brown-
P. nigra subsp. nigra, from which it differs in having ish banded areas. Branches assurgent to irregularly
reddish bark on the branches and nearly flat apo- disposed, forming a conical crown with assurgent
physes on the seed cone scales. From P. sylvestris it branches in young trees and a broad, irregular, very
720 differs in having longer, thicker, dark green leaves open crown in mature trees. Shoots elongating be-
with medial resin ducts and nearly sessile cones. fore leaves start to grow, becoming stiff, assurgent,
greyish white, sometimes glaucous. Cataphylls 5–8
mm long, subulate, brittle, with erose margins,
Taxonomic notes
dark brown. Vegetative buds ovoid-acute to oval;
This is perhaps the better known hybrid taxon in- terminal bud ca. 10 mm long; lateral buds smaller,
volving Scots pine and Black pine Pinus sylvestris resinous. Fascicle sheaths initially 7–9 mm long,
L. × P. nigra J. F. Arnold; other hybrids between the persistent, dark brown, weathering grey. Leaves in
two have been described and named. Controlled fascicles of 3, rarely 4, connate, appearing as a single
­hybridization between these species has also been leaf, not parting until the final year, persisting 2–3
attempted and reported to be successful (Vidaković, years, stiff, straight or slightly curved, often twisted,
1991), so it is probable that hybrids occur occasion- 4–8(–10) cm long, 0.7–0.8 mm wide, with serrulate
ally in nature where the two species meet, as in margins and acuminate apex, dark green. Stomata
­Austria. on all faces of leaves. Pollen cones forming an elon-
gated ‘spike’, ovoid-oblong to cylindrical, 7–9 × 3–3.5
mm, pinkish, turning brown. Seed cones solitary
Distribution
or in pairs on 2.5–6.5 cm long, recurved peduncles.
Austria (limited to the more restricted distribution Mature cones irregularly cylindrical, usually fall-
of P. nigra subsp. nigra and vicinity). ing from the persistent peduncle, (5–)7–12 × 4–5.5
TDWG codes: 11 AUT-AU cm when open. Seed scales parting slightly or more
widely but usually not enough to allow the seeds to
fall, rather weakly attached to a thick rachis with a
Conservation
cuneate base (thereby moveable), thick, irregular,
IUCN: LC with 1–2 deep, cup-like depressions holding the
seeds. Apophysis irregular, prominently raised and
transversely keeled, often rugose, red-brown to dark
Pinus nelsonii Shaw, Gard. Chron., ser. 3, 36: brown. Umbo dorsal, transversely keeled, obtuse, up
122. 1904 [“nelsoni”]. Type: Mexico: Tamaulipas, to 3–4 mm high, with a small, triangular spine. Seeds
Miquihuana, E. W. Nelson 4501 (holotype US). Fig. obliquely obovoid, 12–15 × 8–10 mm, the integument
235 hard, 1 mm thick, pale to dark brown; wing absent.

Etymology Taxonomic notes

This species was named after E. W. Nelson, who col- Pinus nelsonii is a fairly unique species in subgenus
lected the type specimen in June 1898. Strobus, section Parrya. Its position varies somewhat
with the data and other taxa used in cladistic analy-
ses (molecular as well as morphological data), but
Vernacular names
is probably basal in the section, or perhaps sister to
Nelson Pinyon pine, Nelson’s pine; piñon prieto an expanded group of ‘pinyon pines’ (see Phylogeny
(Mexico) and classification of the genus Pinus in Farjon, 2005b:
218–224). This species shares various ­morphological
characters with Asian ‘lace-bark’ pines and with the IUCN: VU (C2a)
Mexican species P. pinceana. However, based on a
wider sampling of data and species as in the above
Uses
mentioned analyses, these taxa do not appear to be
more closely related to P. nelsonii than to other spe- There is no commercial exploitation of this species
cies in the section or even in the entire subgenus due to its low stature and its rarity in remote and
Strobus. inaccessible locations. Its seeds resemble those of
the true pinyon pines (Pinus subsect. Cembroides)
and are edible like these, but the seed crop is usually
Distribution
low compared with these species. Apart from a few
Mexico: Coahuila (Mont. del Carmen), Nuevo León, specimens in botanic gardens and other dendrologi- 721
San Luis Potosí, Tamaulipas. cal collections, this interesting pine is not known in
TDWG codes: 79 MXE-CO MXE-NL MXE-SL MXE- horticulture. It should be suitable to Mediterranean-
TA type climate regions and other dry, warm temper-
ate areas in Europe, the USA, Australia, and South
Africa.
Ecology
Pinus nelsonii is a rare pine occurring in the semi-­
arid foothills and on mesas of the Sierra Madre Pinus nigra J. F. Arnold, Reise Mariazell: 8, t. s.n.
Oriental; the most extensive populations are found 1785.
around the Sierra Peña Nevada in Nuevo León-Tam-
aulipas. It is restricted to sites on rocky limestone
Etymology
with shallow soils. Its altitudinal range is 1600–
2300(–2450) m a.s.l. Annual precipitation ranges The species epithet means black, this probably refers
from 300–600 mm, falling mostly in the summer to the bark which is much darker than that of P. syl­
during brief thunderstorms. Associated conifers are vestris, with which it is often found together.
P. cembroides, P. remota and Juniperus spp. It occurs
in a scrubland zone with deciduous woody taxa, e.g.
Vernacular names
Quercus, Mahonia, ­Comarostaphylis, ­Brahea, and
Sophora, and arborescent monocots, such as Yucca Austrian pine, Black pine; pino nigro (Spanish); pin
and Dasi­lyrion. At higher altitudes it may grade into noir (French); Schwarzföhre (German); karaçam
Pinyon-Juniper woodland, while lower down it is (Turkish)
bounded by a hotter and drier semi-desert scrubland
often dominated by Cactaceae and Yucca spp. Like
Description
several other narrow endemic conifers, P. ­nelsonii is
probably an edaphic relict on limestone. Trees to 40 m tall (–50 m in Corsica), max. d.b.h.
1.89 m (a tree in Corsica), monopodial but some-
times with multiple stems from a low basal trunk.
Conservation
Bark becoming thick and breaking into scaly ridges
Pinus nelsonii is by all accounts a rare pine with a or large plates, separated by irregular, dark fissures,
scattered occurrence largely limited to limestone grey with pinkish or purplish hue, dark grey-brown
outcrops. Its total ‘population’ almost certainly or nearly black. Branches spreading and ascending,
numbers fewer than 10,000 mature trees, mostly in sometimes heavy, forming broadly conical to domed
(sub)populations of a few hundred individuals. It is crowns. Foliage branches stout, rough with pulvini
thought to decline due to habitat disturbance and from fallen leaf fascicles, glabrous, new shoots yel-
loss associated with increased cattle ranging and lowish green, becoming light orange-brown to red-
incidence of destructive fires associated with this brown. Buds ovoid to oblong-conical, sharply acute,
type of land use. Several localities with this little tree non-resinous or resinous; cataphylls brown, thin,
reported in the literature or with older herbarium with papery, grey fringes. Leaves in ­fascicles of 2,
­collections have not been retraced in recent years held by a persistent, 10–12 mm long basal sheath,
and the populations may have disappeared. remaining 2–3 years on branchlets, straight or more
 often curved, rigid or flexible, (4–)8–16(–18) cm along the Black Sea coast eastwards to Krasnodar in
long, sometimes twisted, 1–2 mm wide, light green the Caucasus.
or dark green; margins minutely serrate; apex acute; TDWG codes: 11 AUT-AU 12 COR FRA-FR SPA-AN
stomata in fine lines on all faces. Pollen cones clus- SPA-SP 13 ALB BUL GRC ITA-IT KRI ROM SIC-SI TUE
tered near base of new shoots, spirally arranged, YUG-BH YUG-CR YUG-KO YUG-MA YUG-MN YUG-
ovoid-conical to short cylindrical, 1.5–2.5 cm long, SE YUG-SL 14 KRY 20 ALG MOR-MO 33 NCS-KR 34
0.5–0.7 mm wide, yellow. Seed cones solitary or in CYP EAI TUR
whorls of 2–5 on short peduncles, falling shortly af-
ter seed dispersal, ovoid-conical when closed, (3.5–)
Ecology
5–10(–12) cm long, 2–4 cm wide, opening to ovoid
722 or slightly curved ovoid, usually light brown or pale The wide but fragmented (disjunct) range of P. ­nigra
grey-brown, rarely darker. Seed scales thin woody, throughout S Europe and Turkey guarantees a di-
rigid, narrowly oblong; apophyses slightly raised, verse ecology. It is generally a lower montane spe-
transversely keeled, light yellowish brown, more or cies, but around the Black Sea it is found in hills. It
less lustrous; umbo dorsal, small, usually unarmed grows on a variety of soils, from podzolic sands to
or with a minute, deciduous prickle. Seeds obovoid, limestone, often dependent on region and climate. It
slightly flattened, (4–)6–8 mm long, grey or mottled can form pure stands (which may have been helped
darker; wing obliquely oblong, 15–25 mm long, pale by foresters), but is more commonly associated with
light brown. Pinus sylvestris throughout its range, while region-
ally P. halepensis, P. brutia, P. mugo, P. pinea, P. peuce,
or P. heldreichii can be found with it. It is more tol-
Taxonomic notes
erant to maritime influences like salt-laden winds
Few other species have a taxonomic history as con- than P. sylvestris, so it often occurs closer to the sea.
voluted as Pinus nigra, with numerous taxa at all The geographic variation is partly ecologically de-
ranks from species down to forma decribed and termined, with subsp. laricio more salt tolerant than
named, often in a three tiered system with varieties subsp. nigra, which occurs further inland. Under-
within subspecies within species (for a fairly com- growth in dense pine forests dominated by this spe-
prehensive list of names and their synonyms with cies is usually sparse; more often it forms a mozaic
references see Farjon, 1998, [2001]). It is almost im- with heathland (Erica, Calluna, Vaccinium), which
possible to list all the synonyms as many were pub- can also be present under more open canopies after
lished in regional Floras or periodicals that have not selective felling or natural disturbance. The exten-
even reached the major botanical libraries. Especial- sive plantations and forest management of this spe-
ly in southern and southeastern Europe a tendency cies in Europe over several centuries have made the
to split this species on perceived differences of all distinctions with natural forests less clear.
kinds, including numbers of resin ducts in needles,
is still prevalent (see e.g. Vidacović, 1991). While it
Uses
is appropriate to recognize a few infraspecific taxa
in a species with a variable morphology and a wide Austrian or Black pine is an important timber tree as
but disjunct distribution, extreme splitting would well as amenity tree and has been extensively planted
amount to a refutation that a species after all is made in Europe and to a lesser extent in the USA. In Aus-
up of variable individuals and ditto populations. tria and the Balkans, its wood is traditionally used
A conservative treatment is favoured here, largely to build houses; modern uses include interior floor-
based on the treatment in Flora Europaea 1 (1993). ing (the stage of the Viennese State Opera House is
Even among the five subspecies recognized here, the made of this wood!) and paneling, doors, staircases,
distinctions are often not as clear-cut as is claimed furniture, etc. In the past there was a substantial in-
by those who are thoroughly convinced of them. dustry based on resin tapping, but this has almost
disappeared. In the Mediterranean, it is the major
pine for general construction, fuelwood, pulp for pa-
Distribution
per and to make crates and pallets. Along the North
SW, S and SE Europe; N Algeria; N Morocco; Cy- Sea coast it has been used to stabilize coastal dunes,
prus; Turkey; from the Krym [Crimea] in Ukraine especially subsp. laricio (Corsican pine) which is the
most salt wind tolerant form of the species. The sub- dermal layers and 10–11 resin ducts; seed cones
species pallasiana (Crimean pine), native in Turkey small, (3–)4–6(–7) cm long.
and around the Black Sea, often grows with multi-
ple stems above a basal trunk when in cultivation in
Distribution
NW Europe and has attractive bark with large, light
grey plates, so it was extensively planted as speci- SE Europe: Croatia.
men trees in Victorian period gardens and parks in TDWG codes: 13 YUG-CR
Britain. The species is also tolerant to various forms
of industrial pollution (especially subsp. nigra, Aus-
Conservation
trian pine), and in the USA in particular it is there-
fore popular for plantings in urban and industrial This subspecies of Pinus nigra has a limited distri- 723
settings. There are a number of cultivars named and bution along the coast and on some islands in the
in the horticultural trade. Adriatic Sea and is mainly threatened by habitat
degradation. Many areas are seriously overgrazed
5 subspecies and 1 variety are recognized:
especially by goats, preventing regeneration.
IUCN: VU (B1+2c)
Pinus nigra J. F. Arnold subsp. nigra. Type: Illustra-
tion in J. F. Arnold, Reise Mariazell: 8, t. s.n. 1785.
Pinus nigra J. F. Arnold subsp. laricio (Poir.) Maire,
Pinus nigra J. F. Arnold subsp. croatica Lovric, Oes- Bull. Soc. Hist. Nat. Afrique N. 19: 66. 1928. Pinus
terr. Bot. Z. 119: 569. 1972. laricio Poir., in Lamarck, Encycl. 5: 839. 1804, non
Savi (1798. Type not designated. Fig. 236
Description
Pinus laricio Poir. var. calabrica Loudon, Arbor. Frut.
Trees to 40 m tall; bark breaking into longitudinal Brit. 4: 2201. 1838; Pinus nigra J. F. Arnold var. calabri­
ridges and fissures and small, scaly plates, dark grey. ca (Loudon) C. K. Schneid., in Silva-Tarouca, Uns.
Leaves rigid, curved, 8–13 cm long, dark green, with Freil.-Nadelhölzer: 261. 1913; Pinus nigra J. F. Arnold
2–3 hypodermal cell layers. subsp. calabrica (Loudon) E. Murray, Kalmia 13: 23.
1983; Pinus laricio Poir. subsp. calabrica (Loudon)
Cesca & Peruzzi, Caryologia 55 (1): 24. 2002.
Distribution
E Austria; Balkan Peninsula; Bulgaria; Romania;
Description
Turkey-in-Europe.
TDWG codes: 11 AUT-AU 13 ALB BUL GRC ROM Trees to 50 m tall; bark grey, with deep, longitudinal
TUE YUG-BH YUG-CR YUG-KO YUG-MA YUG-MN fissures and irregular, scaly plates. Foliage branches
YUG-SE YUG-SL orange-brown, becoming light grey-brown; leaves
8–15 cm long, slender, flexible, 1.2–1.5 mm wide,
light green, with 1–2 hypodermal cell layers.
Conservation
IUCN: LC
Distribution
Pinus nigra J. F. Arnold subsp. dalmatica (Vis.) Mediterranean: Corsica, Italy.
Franco, Dendrol. Florest.: 55. 1943. Pinus dalmatica TDWG codes: 12 COR 13 ITA-IT
Vis., Fl. dalm. 1: 199. 1842. [in obs.]; Fl. dalm. suppl.:
43. 1872; Pinus nigra J. F. Arnold var. dalmatica
Conservation
(Vis.) Businský, Acta Pruhon. 88: 11. 2008. Type not
designated. IUCN: LC

Description
Small trees to 15 m tall, often with flat-topped
crowns; leaves short, 4–7 cm long, with 2–5 hypo­
 Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
Taxonomic notes
Holmboe var. pallasiana, [Stud. Veg. Cyprus]
Bergens Mus. Skr., ser. 2, 1 (2): 29. 1914. Pinus This apparent mutational form would have been
pallasiana Lamb., Descr. Pinus 2: 1, t. 1. 1824. Type: more appropriately treated as a forma, not a variety.
Illustration in Lambert, Descr. Pinus 2: t. 1. 1824 It was earlier named as Pinus nigra var. pyramidata,
(lectotype, designated here). a later homonym, hence the new name published by
Businský (op. cit.).
Pinus laricio Poir. var. caramanica Loudon, Arbor.
Frut. Brit. 4: 2201. 1838; Pinus nigra J. F. Arnold var.
Distribution
caramanica (Loudon) Rehd., Man. Cult. Trees: 61.
724 1927; Pinus nigra J. F. Arnold subsp. caramanica Turkey: Kütahya, near Dulkadir village.
(Loudon) Businský, Acta Pruhoniciana 68: 22. 1999;
Pinus pallasiana Lamb. subsp. caramanica (Lou- Pinus nigra J. F. Arnold subsp. salzmannii (Dunal)
don) Chrtek & B. Slavik, Flora Mediterranea 10: 236. Franco, Dendrol. Florest.: 56. 1943. Pinus salz­
2000. mannii Dunal, Mém. Sect. Sci. Acad. Sci. Mont­
Pinus nigra J. F. Arnold var. yaltirikiana C. U. Alpte­ pellier 2: 82. 1851; Pinus nigra J. F. Arnold var.
kin, Istanbul Univ. Orman Fak. Dergisi, ser. A, 36 salzmannii (Dunal) Laguna Lumbreras, Flora
(2): 147. 1987. Montiberica 15: 28. 2000. Type: France: Languedoc,
Pinus nigra J. F. Arnold var. columnaris-pendula Hérault, St. Guillaume-le-désert, P. Salzmann s.n.
Boydak, Karaca Arbor. Mag. 6 (1): 17. 2001. (holotype MPU). Fig. 237

Pinus clusiana Clemente, in Arias, Adic. Agric. Gen.

Description
Herrera 2: 404. 1818; Pinus nigra J. F. Arnold subsp.
Trees to 40 m tall, commonly (in cultivation?) clusiana (Clemente) Rivas-Martínez, Mem. Mapa
forked into 2 or more ascending trunks; bark light Ser. Veget. Potencial España: 146. 1988.
grey, with deep, longitudinal, dark fissures and often Pinus laricio Poir. var. angustisquama Willk., in
very large, irregular, scaly plates. Foliage branches Willkomm & Lange, Prodr. Fl. Hispan. 1: 18, 73. 1870;
orange-brown; leaves rigid, straight or curved, (8– Pinus nigra J. F. Arnold var. angustisquama (Willk.)
)12–17 cm long, light green, with 2–5 hypodermal Laguna Lumbreras, Flora Montiberica 15: 28. 2000.
cell layers and 6–9 resin ducts. Pinus laricio Poir. var. latisquama Willk., in
Willkomm & Lange, Prodr. Fl. Hispan. 1: 18, 73. 1870;
Pinus nigra J. F. Arnold var. latisquama (Willk.) La-
Distribution
guna Lumbreras, Flora Montiberica 15: 28. 2000.
Cyprus; from the Krym [Crimea] along the Black
Sea coast to Krasnodar; Turkey.
Description
TDWG codes: 14 KRY 33 NCS-KR 34 CYP TUR
Trees to 30 m tall; bark with irregular, scaly plates,
variously fissured, light brown to grey-brown. Buds
Conservation
resinous; leaves slender, 8–16 cm long, 1–1.3 mm
IUCN: LC wide, flexible, with (1–)2 hypodermal cell layers.

Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)

Distribution
Holmboe var. fastigiata Businský, Acta Pruhon. 88:
8. 2008. SW Europe, France (Cevennes, Pyrenees), Spain;
Algeria (Montagnes du Hodna); Morocco (Rif
Mountains).
Description
TDWG codes: 12 FRA-FR SPA-AN SPA-SP 20 ALG
A fastigiate branching growth form of Pinus nigra MOR-MO
subsp. pallasiana.
 slightly raised, transversely keeled, rhombic to pen-
Conservation
tagonal in outline, on the proximal scales more or
IUCN: LC less gibbous, (lustrous) dark brown, radially striate,
weathering dull grey. Umbo dorsal, raised and of-
ten curved, usually armed with an inflexed, 2–3 mm
Pinus occidentalis Sw., Prodr.: 103. 1788. Type: long spine. Seeds obliquely obovoid, flattened, 5–6
Haiti: Nippes, [“Hispaniola: aux Pins, quartier des × 3–4 mm, light, mottled grey-brown. Seed wings
Nippes, du côté septentrional”], O. P. Swartz s.n. obliquely ovate or oblong, 12–18 × 4–6 mm, ochra-
(lectotype BM). ceous with black or grey tinge or stripes.

Pinus occidentalis Sw. var. baorucoensis Silba, Phyto- 725

Distribution
logia 58: 368.
Hispaniola: Dominican Republic; Haiti.
TDWG codes: 81 DOM HAI-HA
Etymology
The species epithet means ‘from the west’ (where it
Ecology
occurs from a European perspective).
This species occurs in diverse habitats from the low-
lands at about 200 m a.s.l. to the highest mountain
Vernacular names
ridges (Pico Duarte and Pico La Pelona) on the is-
Cuban pine is a misnomer, it should be more appro- land at almost 3200 m. The more extensive and pure
priately named Hispaniolan pine. stands occur from 900–2700 m, but in more acces-
sible areas these are much depleted. Soils are either
derived from limestone at lower altitudes, or more
Description
acid, clay-like and shallow in the Cordillera Central.
Trees to 30–40 m tall, d.b.h. to 1–1.2 m; trunk mo- Pinus occidentalis consequently is found in a variety
nopodial, erect. Bark thick, rough, scaly, break- of vegetation types, mostly occupying the shallow,
ing into irregularly square plates divided by deep nutrient-poor soils and rock outcrops, where it may
fissures, turning grey-brown to grey. Branches occur in open or dense, pure stands or mixed with
spreading or ascending, in lower crown curved and various broad-leaved trees and shrubs. In disturbed
drooping, forming an irregular, open crown. Shoots (grazed) areas Pteridium aquilinum can dominate
uni-nodal, glaucous to pruinose in the first year, the ground cover; in frequently burnt areas grasses
later brown. Cataphylls slender, subulate to caudate, (e.g. Danthonia domingensis, Andropogon spp.) and
straight or recurved, scarious, brown. Vegetative again Pteridium replace shrubs and small trees. An-
buds ovoid to ovoid-oblong, acute; terminal bud nual precipitation varies greatly with exposition, but
10–15 mm long; lateral buds shorter, usually slight- ranges between 1200–1600 mm where most pine
ly resinous. Fascicle sheaths persistent, (8–)10–15 forests occur, it exceeds 2300 mm in the N and E of
mm long, lustrous silvery brown, weathering grey- the Cordillera Central. There is a 3–5 month dry sea-
brown. Leaves in fascicles of 3–5, persisting to third son during winter, which may bring frost, but rarely
year, straight or slightly curved, more or less rigid, snow, at the higher altitudes above 1600–1800 m.
(11–)14–18(–20) cm long, 1.2–1.4 mm wide, with
serrulate margins, acute, pungent, light green. Sto-
Conservation
mata on all faces of leaves. Pollen cones cylindrical,
1.5–2.5 cm × 5 mm, pinkish yellow, turning yellowish Being the only species of Pinus on Hispaniola and
brown. Seed cones sub-terminal, solitary or in pairs formerly abundant over much of the island, it has
on 1–2 cm long, straight or recurved peduncles, per- been heavily exploited for timber. According to Dar-
sisting several years after seed dispersal, falling with row & Zanoni (1991) it has been depleted from an
peduncle attached. Mature cones ovoid to ovoid- estimated 3 million ha of primeval more or less pure
conical, straight or curved, nearly symmetrical, pine forests to perhaps less than 5% of that area, but
(4–)5–9(–11) × 3.5–6.5 cm when open. Seed scales accurate estimates of even the present forest extent
oblong, straight or recurved, thin woody. Apophysis are lacking. Protection in the Dominican Republic
 is inadequate, but existent, contrary to the virtually i­ rregular, open crown. Shoots rough and scaly, red-
uncontrolled situation in Haiti. dish brown. Cataphylls 10–15 mm long, subulate, re-
IUCN: NT curved, scarious, brown, weathering blackish grey.
Vegetative buds ovoid-oblong to fusiform; terminal
bud 15–25 mm long; lateral buds ovoid-acute, small-
Uses
er, none resinous. Fascicle sheaths up to 25 mm long,
Cuban pine (Hispaniolan pine) is an important persistent, not reduced in length, red-brown weath-
timber tree on its native island Hispaniola, where ering to nearly black. Leaves in fascicles of 4–5, per-
despite intensive exploitation it is still common. Its sisting 2–3 years, straight, rigid or less commonly
wood has good qualities comparable to those found pliant, (11–)14–25(–30) cm long, 0.8–1.6 mm wide,
726 in the more widespread species P. caribaea and is with serrulate margins, acute-pungent, lustrous (yel-
used as round wood for transmission poles, fence lowish) green. Stomata on all faces of leaves. Pollen
posts, construction timber, crates, boxes, and made cones oblong-cylindrical, 1.5–2 cm × 5–6 mm, pink
into wood pulp for particleboard as well as paper. or reddish, turning yellowish brown. Seed cones
There is limited resin tapping for local use only. It subterminal, solitary or in whorls of 2–4 on stout, up
is a tropical pine and even those trees at the highest to 35 mm long, recurved peduncles which eventu-
altitudes in the Dominican Republic are not likely to ally fall with cones, persisting for several years after
yield progeny that can be grown successfully in cool seed dispersal. Mature cones broadly ovoid to sub-
temperate climates. Growing pines in greenhouses globose, semi-serotinous, when opened often wider
is problematic: unlike many other subtropical or than long, with a flattened base, 3–8(–10) × 3–9(–12)
tropical conifers they do not produce strong wood to cm when open. Seed scales thick woody, symmet-
hold the tree upright without the influence of wind rical, oblong, straight or slightly recurved. Apo-
that moves the stems during secondary growth. physis nearly flat or slightly raised, in some cones
pyramidal, (weakly) transversely keeled, rhombic to
pentagonal in outline, lustrous, ochraceous to light
Pinus oocarpa Schiede ex Schltdl., Linnaea 12: brown, weathering grey. Umbo dorsal, flat or raised,
491. 1838. Type: Mexico: Michoacan, Aguililla, Las sometimes curved, obtuse or rarely with a minute
Playitas Forest Station, along the track to Cañas, prickle. Seeds obliquely ovoid, slightly flattened, 4–8
B. T. Styles 36 (neotype FHO). × 3–4.5 mm, blackish grey, often with black spots.
Seed wings 8–18 × 4–8 mm, greyish brown.
Pinus oocarpa Schiede ex Schltdl. var. manzanoi
Martínez, Anales Inst. Biol. Univ. Nac. México 11:
Distribution
70. 1940.
Mexico: from the Sierra Madre Occidental SE to
Mesoamerica, in S Sonora, Sinaloa, SW Durango,
Etymology
Nayarit, S Zacatecas, Jalisco, Michoacán, México,
The species epithet (Greek: oos = egg, carpos = fruit) Distrito Federal, Hidalgo, N Puebla, Morelos, Tlax-
refers to the shape of closed seed cones. cala, Guerrero, Oaxaca, S Veracruz and Chiapas;
widespread in the highlands of Guatemala; Hondu-
ras; El Salvador and NW Nicaragua.
Vernacular names
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
Egg-cone pine; ocote chino, pino chino, pino colo- PU MXC-TL MXE-AG MXE-CO MXE-CU MXE-DU
rado (Spanish) MXE-GU MXE-HI MXE-NL MXE-QU MXE-SL MXE-
TA MXE-ZA MXG-VC MXN-SI MXN-SO MXS-CL
MXS-GR MXS-JA MXS-MI MXS-NA MXS-OA MXT-CI
Description
80 ELS GUA HON NIC
Trees to 30–35 m tall, d.b.h. to 1–1.3 m; trunk mo-
nopodial, erect. Bark thick, rough, scaly, breaking
Ecology
into small or large, longitudinal plates and shallow
fissures, dark brown to grey-brown. Branches long, This species extends over a NW-SE distance of ca.
often tortuous, spreading, forming a rounded or 3000 km and consequently is found under very
­ ifferent ecological conditions. This is expressed in
d ­ orticulture it is very rare and limited to a few bo-
h
its altitudinal range, from (200–)500–2300(–2700) tanic gardens and other tree collections in countries
m a.s.l., and in the variation of annual precipita- with warm climates.
tion, from 700–3000 mm. Seasonality is mainly ex-
pressed in a (long) dry season from October to June
in much of its range. Nearly everywhere, fire is an Pinus palustris Mill., Gard. Dict., ed. 8: Pinus No.
integral part of the ecosystem, but man-made fires, 14. 1768. Type not designated.
often deliberate, add substantially to the frequency
with which they occur. Pinus oocarpa has semi-
Etymology
serotinous cones and is adapted to fire, at least at
natural frequencies. It occurs in usually open wood- The species epithet means ‘of the marsh’, referring to 727
land or forest, often in pure stands or as a constitu- the habitat of the species.
ent of pine-oak woodland. Other pines commonly
associated with it are P. engelmannii, P. leiophylla
Vernacular names
and P. douglasiana in the NW and P. maximinoi,
P. devoniana and P. tecunumanii to the SE. If fires are Longleaf pine, Florida pine, Georgia pine
less frequent, there can be an understorey of shrubs
with, e.g. Calliandra, Acacia, Leucaena, Hybosema,
Description
Byrsonima, and Leucothoe, but often the burning fa-
vours Pteridium aquilinum or grasses. Trees to 45(–47) m tall; trunk to 1.2 m d.b.h., bole
straight and columnar. Bark breaking into large, ir-
regularly rectangular, scaly plates separated by deep
Conservation
fissures, orange-brown or reddish brown. Branches
The greatly increased frequency of fires, often delib- relatively sparse, spreading and descending, form-
erately ignited, poses a serious threat to the devel- ing an open, rounded crown. Foliage branches stout,
opment of mature stands of this pine. The problem up to 2 cm thick in leading shoots, upturned at apex,
is especially poignant in Mesoamerica, where both prominently ridged and grooved between decurrent
fires and infestations by the pine bark beetle (Den­ pulvini, glabrous, orange-brown turning darker red-
droctonus mexicanus) are frequently followed by dish brown. Buds large; terminal bud 3–4.5 cm long,
‘lumber salvation operations’ and subsequent con- narrowly ovoid, not resinous; cataphylls narrowly
version into pasture (Perry, 1991). Despite this, the oblong, with fringed margins and recurved at apex,
species is very widespread and numerous in many silvery white. Leaves in fascicles of 3, rarely 2 or 5,
areas, so it does not meet any of IUCN’s criteria of a held together in 20–25(–30) mm long sheaths, per-
threatened species. sisting 2 years, 20–45 cm long (longest in young, vig-
IUCN: LC orous trees), spreading and drooping, slender and
flexible, slightly twisted, ca. 1.5 mm wide; margins
minutely serrulate; apex shortly acute to acuminate;
Uses
leaf colour bright lustrous green; stomata in fine
Throughout its range, Pinus oocarpa is an important lines on all surfaces. Pollen cones clustered at base of
timber tree, especially for local and regional mar- new shoots, spirally arranged, spreading out radially,
kets. The wood is of better quality than that of P. car­ cylindrical, 4–7(–8) cm long, 0.8–1.5 cm wide, pur-
ibaea, it is stronger and less resinous. It is much used plish red, turning red-brown. Seed cones solitary or
for sawn timber, especially applied to light construc- in pairs near ends of foliage branches, sessile or very
tion like carpentry and joinery and floors in hous- short pedunculate, soon falling after seed dispersal,
es. Courser uses are railway sleepers and beams or often leaving a rosette of basal scales on branch, (15–)
transmission poles. The species is also widely tapped 20–25 cm long, symmetric, narrowly ovoid when
for resin, but only for a few years before felling. This closed, broadly ovoid-cylindric when open, with a
species has been introduced to many subtropical flat base. Seed scales oblong, thin woody, rigid, re-
and tropical countries as a plantation forestry tree, curved near base of cone, elsewhere spreading wide,
it is perhaps most successful in W Africa, South dull brown; apophyses slightly raised, transversely
Africa, and in Brazil, Colombia and Argentina. In keeled, rhombic in outline, dull tan or brown in
 colour; umbo dorsal and central, broadly triangu- growth with a grass stage and many sites originally
lar, obtuse, armed with a small, persistent, recurved occupied by P. palustris are now dominated by other
prickle. Seeds obovoid, slightly flattened, 9–12 mm species. Prevention of fires increases competition
long, pale brown mottled with darker specks; wing from herbs and shrubs as well as other pines, as
30–40 mm long, 10–12 mm wide. Seedlings with a the seedlings of P. palustris do not initially grow in
definite and prolonged ‘grass stage’. height, unlike other pines common on the Coastal
Plains. Controlled burning is therefore necessary for
its successful establishment, but only P. palustris will
Distribution
survive this at a juvenile stage and in managed pine
SE USA, from Virginia to E Texas in the Atlantic and forest this practice would exclude other pine species.
728 Gulf Coastal Plains. IUCN: VU (A1c-e)
TDWG codes: 77 TEX 78 ALA FLA GEO LOU MSI
NCA SCA VRG
Uses
Longleaf pine is an important species providing
Ecology
good quality timber as well as naval stores derived
Pinus palustris grows mostly in the warm temperate from its resin. It was heavily exploited since Europe-
to subtropical coastal plain of the SE United States, ans settled in the Coastal Plains and served a major
but it extends into the uplands and foothills of the forest industry in the region. Its wood is used for
southern Appalachian Mountains to about 700 m sawlogs, stage flooring, plywood, pulpwood and
a.s.l. It requires a warm, humid climate and a moist produces poles, fence posts, and piling as it makes
soil, although the latter can vary from wet, poorly straight stems largely free of branches when grown
drained clay in swampland to thin stony soils on in closed stands. Turpentine and other chemicals
rocky mountain slopes and ridges. For the most part can be distilled from the chipped wood and even
soils are sandy, acidic and poor in nutrients. This stumps of trees are pulled from the soil for this pur-
species can grow in pure stands but is often found pose. Longleaf pine is not planted for forestry out-
together with other pines, like P. elliottii, P. echinata side its natural range and it is relatively rare as an
and P. taeda. Many broad-leaved trees (angiosperms) amenity tree, mainly confined to coastal areas in the
grow with it, too, forming mixed forests on mesic SE of the USA. The species is somewhat difficult to
sites with e.g. Quercus spp., Nyssa sylvatica, Liquid­ grow through its ‘grass stage’ to the sapling stage and
ambar styraciflua, Cornus florida, Sassafras albidum, beyond and needs a protected location in a mild cli-
and Diospyros virginiana and many shrubs. In the mate.
foothills in Alabama many oaks (Quercus spp.) ac-
company Pinus palustris, forming mixed pine-oak
woods. This species of pine is well adapted to fires Pinus parviflora Siebold & Zucc., Fl. Japon. 2 (3):
through its grass stage and succeeds quickly from 27, t. 115. 1842.
seed to take advantage of freed space and nutrients,
while it can survive the next ground fire by resprout-
Etymology
ing from the very short stem until it has built enough
of a root system to accelerate its stem growth dra- The species epithet means ‘with little flowers’ which
matically and rise above the damaging flames. perhaps refers to an early stage of the seed cones.

Conservation Vernacular names

Pinus palustris was once the most common pine in Japanese white pine; Goyō-matsu, Hime-komatsu
the Coastal Plains, perhaps covering 25 million ha. (Japanese)
Exploitation for timber and naval stores and conver-
sion to farmland and pasture reduced this by 1985
Description
to less than 1.6 million ha. This decline, although
its rate has slowed, is ongoing. Foresters and land- Trees to 15–20 m tall; trunk to 1 m d.b.h., usually
owners prefer other species that do not delay initial a straight, columnar bole. Bark smooth in young
trees and in upper part of the crown, becoming of growing up under the parent trees. Persistence of
rough and scaly on trunk, shallowly grooved, grey cones may also be part of this syndrome, depending
to dark grey. Branches whorled in young trees, later on the feeding habits of the bird species that feed on
spreading and ascending; stem sometimes forked them and (presumably) play a role in their dispersal.
near the top, forming a conical to rounded crown,
on exposed sites often flattened and open. Foliage
Distribution
branches slender, new shoots initially pubescent,
soon glabrous or with remnant hairs in grooves, Japan: Hokkaido, Honshu, Kyushu, Shikoku; South
grey-green turning light brown. Buds small, ovoid, Korea, Utsurio-To (island).
non-resinous or slightly resinous; cataphylls free at TDWG codes: 38 JAP-HK JAP-HN JAP-KY JAP-SH
apex, light orange-brown. Leaves crowded in dense KOR-SK 729
tufts towards end of shoots, persisting 3–5 years, in
fascicles of 5, held by deciduous sheaths with thin
Ecology
scales falling in the second year, curved and slightly
twisted, (3–)4–7(–8) cm long, slender and flexible, Pinus parviflora occurs at altitudes from just above
0.7–1 mm wide, triangular in cross-section, often sea level to about 2500 m, with an optimum between
twisted, green to glaucous green; stomata in 3–4 1000 and 1500 m a.s.l. in montane forests. It is found
prominent white lines on the two adaxial surfaces; in both pure and mixed stands with other conifers
margins minutely serrulate; apex acute. Pollen cones and also with angiosperms, usually on steep slopes,
in small clusters near base of new shoots, spirally ar- dry sites or rocky ridges. At high, subalpine habi-
ranged, ovoid-oblong to cylindrical, 1.5–2.5 cm long, tats this species becomes dwarfed and some of the
yellow turning light brown. Seed cones in whorls cultivated forms may have been derived from such
of 4–8(–10) on branches, persistent, sessile or very provenances.
short pedunculate, spreading, when still growing
ovoid or subglobose, green or glaucous, very resin-
Uses
ous, with opened scales 4–7 cm long and to 4 cm
wide, becoming more ovoid or irregular. Seed scales Japanese white pine has little value as a timber tree,
few, thin woody, flexible, with larger scales straight but great value as an ornamental. In Japan, this spe-
or curved, but basal smaller scales not recurved; ap- cies has been in cultivation in gardens and parks
ophyses variously shaped, strongly concavo-convex and in temple grounds for many centuries. Growth
(‘cupped’) or more or less flat, incurved, becoming forms that stay low and form picturesque shapes are
dull brown; umbo terminal, obtuse. Seeds ellipsoid- highly sought after and these preferences culminate
ovoid or obovoid, 7–10 mm long, 5–6 mm wide; in the art of bonsai, for which this is the most com-
wing to 12 mm long, or rudimentary. monly used species of pine. In bonsai culture growth
is inhibited and plants are slowly forced by various
techniques into shapes that appear to imitate ancient
Taxonomic notes
trees growing on exposed mountains in a miniature
Pinus parviflora is closely related to a group of pines fashion. In Western horticulture, the species is also
that includes P. fenzeliana (syn. P. kwangtungensis), popular and a substantial number of cultivars has
P. dalatensis, P. wangii, and P. morrisonicola, but oc- been produced, predominantly dwarf forms with
curs at a considerable distance from these in Japan. strongly glaucous foliage and some with variegated
One variety is commonly recognized: P. parviflora needles. Naturally growing trees of the species are
var. pentaphylla (Mayr) A. Henry (syn. P. penta­ relatively rare and mostly confined to collections in
phylla Mayr), with a more or less well developed arboreta. Most plants in the trade under the species
seed wing and flatter, less cup-like seed cone scales. name are derived from Japanese garden selections
Wingless seeds and hollowed, cup-like scales that (cultivars without a name).
accommodate them are not independent characters
but form an ‘adaptive complex’ to prevent the seeds 2 varieties are recognized:
from falling where the seedlings have little chance
 Pinus parviflora Siebold & Zucc. var. parviflora.
Vernacular names
Type: Japan: [locality unknown], “Pinus cembra”,
P. F. von Siebold s.n. (lectotype L, acc. no. 901.138– Jelecote pine, Spreading-leaved pine, Mexican weep-
394). ing pine; ocote, pino triste (Mexico)

Description Description
Seed scales cup-like in shape, with deep seed cavi- Trees to 35–40 m tall, d.b.h. to 1 m; trunk monopo-
ties. Wings of seeds 3–7 mm long, 6–8 mm wide, ru- dial, erect, straight. Bark thick, rough and scaly,
dimentary (shorter than the seed body). with large elongated plates and deep, longitudi-
730 nal fissures, dark grey-brown below, lighter above.
Branches spreading or slightly ascending; branches
Distribution
of higher orders slender, drooping or sub-pendulous,
Japan: Honshu, Kyushu, Shikoku. with ultimate branches pendant, forming a conical
TDWG codes: 38 JAP-HN JAP-KY JAP-SH to domed, open crown. Shoots often multi-nodal,
rough and scaly, yellowish to reddish brown. Cata-
phylls subulate, recurved at apex, scarious, brown.
Conservation
Vegetative buds oval-oblong to cylindrical; terminal
IUCN: LC bud 15–20 mm long; lateral buds shorter, all brown,
not resinous. Fascicle sheaths initially long, 20–30
Pinus parviflora Siebold & Zucc. var. pentaphylla mm, persistent but reduced to 12–15 mm in mature
(Mayr) A. Henry, in Elwes & Henry, Trees Gr. fascicles. Leaves in fascicles of 3–4(–5), persisting
Brit. Ireland 5: 1033. 1910. Pinus pentaphylla Mayr, 2–3 years, thin, lax, drooping to pendant, (11–)15–
Monogr. Abiet. Japan. Reich.: 78. 1890; Pinus parvi­ 25(–30) cm long, 0.7–0.9(–1) mm wide, serrulate at
flora Siebold & Zucc. subsp. pentaphylla (Mayr) margins, acute, pale green to dark green. Stomata on
Businský, Acta Pruhoniciana 68: 12. 1999. Type not all faces of the leaves. Pollen cones ovoid-oblong to
designated. cylindrical, 1.5–2 cm × 5–6 mm, pinkish yellow. Seed
cones subterminal or lateral, in whorls of 2-many,
rarely solitary, persistent or deciduous, pedunculate.
Description
Mature cones ovoid or ovoid-attenuate when closed,
Seed scales more or less flat, not cup-like in shape. usually slightly curved, 5–10(–12) × (3–)4–6.5 cm
Wings of seeds 10–12 mm long, 6–8 mm wide (ap- when open. Seed scales oblong, usually curved
prox. as long as the seed body). when spreading, the proximal scales connate. Ap-
ophysis nearly flat to slightly raised, transversely
keeled, more or less gibbous on the proximal scales,
Distribution
more so on one side of the cone; upper margin acute
Japan: Hokkaido, Honshu; South Korea: Utsurio-To. or obtuse-rounded; colour yellowish brown. Umbo
TDWG codes: 38 JAP-HK JAP-HN KOR-SK dorsal, flat or raised, often sunken into the apophy-
sis, with a minute, deciduous prickle. Seeds 4–6 ×
2–4 mm, blackish grey. Seed wings 12–18 × 5–8 mm,
Conservation
light brown with dark stripes.
IUCN: LC
Distribution
Pinus patula Schiede ex Schltdl. & Cham., Linnaea
Mexico: Tamaulipas, Querétaro, Hidalgo, México,
6: 354. 1831.
Distrito Federal, Morelos, Tlaxcala, Puebla, Ver-
acruz, Oaxaca and Chiapas.
Etymology
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
The species epithet means ‘spreading wide’ and re- PU MXC-TL MXE-HI MXE-QU MXE-TA MXG-VC
fers to the leaves, which are in fact drooping or pen- MXS-OA MXT-CI
dulous.
 Pinus patula Schiede ex Schltdl. & Cham. var.
Ecology
patula. Type: Mexico: Veracruz, Cruz Blanca, [“In-
Pinus patula occurs in the warm to temperate high- ter Cruz blanca & Jalacingo”], C. J. W. Schiede &
lands of central and southern Mexico, at altitudes F. Deppe 1108 (lectotype HAL, sheet 1/4). Fig. 238, 239
between 1500 and 3000 m a.s.l. and in areas with
abundant precipitation ranging from 1000–2200
Description
mm. According to altitude, the climate ranges from
subtropical to temperate. Fog plays a significant Seed cones (very) short pedunculate, seemingly ses-
part in the availability of moisture through much of sile at maturity, in whorls of 2-many, tenacious, per-
the dry season. It occurs in a variety of forest types, sisting many years, serotinous, ovoid when closed,
with conifers or with angiosperms, especially in the broadly ovoid and (5–)7–10(–12) × (3–)4–6.5 cm 731
Liquid­ambar forests on the eastern slopes facing the when open. Seed scales ca. 120–150, (thick) woody,
Caribbean Sea which receive most of the rain and rigid, many of the proximal scales remaining con-
fog. These forests are famous for their abundance nate.
of epiphytes, among which are ferns, bromeliads
and orchids. Pinus patula is also associated with
Distribution
other pines, e.g. Pinus pseudostrobus, P. maximinoi,
P. ­ayacahuite, and locally, P. greggii, in mixed pine or Mexico: in a few localities in Tamaulipas, in Queré-
pine-oak forest; in a few places it is found associated taro, Hidalgo, México, Distrito Federal, Morelos,
with Abies religiosa. Tlaxcala, Puebla, Veracruz, Oaxaca and Chiapas.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
PU MXC-TL MXE-HI MXE-QU MXE-TA MXG-VC
Uses
MXS-OA MXT-CI
This species is one of the most important pines for
timber in Mexico, as it grows fast and produces a
Conservation
long, straight bole, for the most part free of branch-
es. It is also widely introduced in other tropical IUCN: LC
countries in plantation forestry, where in some cases
it has become problematic as an invasive species. In Pinus patula Schiede ex Schltdl. & Cham. var. lon­
southern and eastern Africa it is planted commer- gipedunculata Loock ex Martínez, Pinos Mexic., ed.
cially; other countries where it has been introduced 2: 333, f. 276–280. 1948 [“longepedunculata”]. Pinus
for plantation forestry on a large scale are Colombia, ×longipedunculata (Loock ex Martínez) Businský,
Brazil and Argentina and to a more limited extent Acta Pruhon. 88: 11. 2008. Type: Mexico: Oaxaca,
India, Nepal and New Zealand. Much work has been Sierra Benito Juarez, Rancho Benito Juarez, (Ran-
done by forest geneticists and tree growers in breed- cho Tablas), E. E. M. Loock 113a (holotype PRF).
ing programmes to improve seed provenances for
timber growing in tropical countries. The wood is
Description
soft and light coloured and easily worked, but can
be susceptible to blue stain without treatment. It Seed cones distinctly pedunculate, peduncles up
finds applications in flooring and paneling, plywood to 20 mm long, curved, falling with cones after a
and particleboard manufacture, veneers, crates and few years. Cones opening readily, narrowly ovoid
boxes, and of course for its softness, pulp for paper; to ovoid-attenuate when closed, (narrowly) ovoid
it is therefore less suitable for high quality furniture when open, then 5–8 × 3.5–4.5(–5) cm. Seed scales
and tools. In horticulture it has made some progress ca. 100–120, (thin) woody, somewhat flexible, part-
in recent years as it forms an attractive tree with its ing readily to release the seeds.
fine foliage and several provenances have proved to
be quite hardy.
Distribution
2 varieties are recognized: Mexico: known from only two localities in Hidalgo
and Veracruz, but more widespread in Oaxaca, also
in central Chiapas.
 TDWG codes: 79 MXE-HI MXG-VC MXS-OA MXT- 2–3(–4) cm wide when closed, opening to 4–5(–6)
CI cm wide, resinous. Seed scales thin woody, flexible,
cuneate to obovate, more or less flat; basal scales not
recurved but imbricate; apophyses broadly rhom-
Conservation
bic to rounded distally, slightly thickened, mostly
IUCN: LC incurved at apex, longitudinally striate or grooved,
light brown when dried; umbo terminal, obtuse,
dark grey-brown. Seeds obovoid, slightly flattened,
Pinus peuce Griseb., Spicil. Fl. Rumel. 2: 349. 1846. (5–)6–8 mm long, grey; wing adnate, 15–20 mm long.
Type: Macedonia: NW Macedonia, Sar Planina,
732 [“in m. Peristeri supra Bitoliam”], A. H. R. Grisebach
Taxonomic notes
s.n. (holotype not located, isotype K).
The relationships of this species, the only European
member of Subsection Strobi in Europe according to
Etymology
the classification adopted in the first edition of the
The species epithet derives from the Greek peuke = book Pines (Farjon, 1984: 194), have recently been
pine tree. revised on the basis of molecular evidence (Liston et
al. in Mill, 2003). The division into subsections Cem­
brae and Strobi based on cone and seed morphology
Vernacular names
appear to break down, with similar results from both
Balkan pine, Macedonian pine chloroplast and nuclear DNA. The wingless seeds,
dispersed by birds, have resulted in a specific type of
cone (see e.g. P. albicaulis and P. cembra) amounting
Description
to convergent evolution. These pines are not neces-
Trees to 25(–30) m tall; trunk to 1 m d.b.h., bole usu- sarily closely related because of these similarities.
ally straight. Bark thin and smooth on young trees Instead, P. cembra is probably more closely related to
and branches, becoming thicker and breaking into P. peuce than to P. albicaulis. A more careful second
small plates, at lower trunk longitudinally fissured, look at anatomy and morphology, avoiding strongly
grey-brown; inner bark purplish brown. Branches adaptive traits that have led to convergence, may
spreading more or less horizontally, relatively short yet detect characters that support the results from
or longer and assurgent in solitary trees, poorly DNA studies, or refute them, as the case may be. It is
self-pruning, forming a dense, conical or narrowly important to include all species of section Quinque­
conical, sometimes broadly conical crown. Foliage folius and appropriate outgroup taxa in such studies.
branches stout; new shoots glabrous, smooth, often
glaucous, turning grey-green and finally grey-brown.
Distribution
Buds ovoid-conical, ca. 10 mm long, acute, resinous
or without resin; cataphylls brown with a narrow, SE Europe: Albania; W Bulgaria; Macedonia; N
whitish margin. Leaves in fascicles of 5, initially held Greece; Serbia.
in a deciduous, 15–18 mm long basal sheath, persist- TDWG codes: 13 ALB GRC YUG-MA YUG-SE
ing 3–4 years in tufts on branchlets, straight, more
or less rigid, 7–10(–12) cm long, slender, 0.5–0.7 mm
Ecology
wide; margins minutely serrulate; leaf colour grey-
green or slightly glaucous; apex acute; stomata in Pinus peuce is a montane pine, growing between 600
fine whitish lines on all surfaces, but only a few on m and 2200 m above sea level and usually on silicate
the abaxial side. Pollen cones few and small, 10–15 rock types. In Albania and Serbia it is also found on
mm long, clustered in spiral arrangement at base of serpentine. It can grow on a variety of soil types, usu-
new shoots, yellow with a purple hue. Seed cones ally poor in nutrients and acidic to basic. In Grecian
solitary or in whorls of 2–4 on short, curved pedun- Macedonia it forms pure stands on gentle mountain
cles, initially erect, green becoming red or purple, slopes, interspersed with grassy glades and mead-
when fully grown becoming pendulous, cylindrical, ows. In most areas where it occurs, it is mixed with
(7–)8–15(–20) cm long, obliquely curved at base, Picea abies and/or Abies alba / A. ×borisii-regis, with
which it can compete due to relatively high shade
Vernacular names
tolerance.
Maritime pine; pino gallego, pino resinero (Span-
ish); pin maritime, pin des landes, pinastre (French);
Conservation
pinastro, pino marittimo (Italian); pinheiro-bravo
This species is probably a Tertiary relict that has sur- (Portuguese)
vived severe contractions of its range due to Alpine
glaciations during the Pleistocene. Its current range
Description
consists of two disjunct populations, one in the west
centered in Albania and one in the east in W Bul- Trees to (25–)30(–40) m tall; trunk to 1.5 m d.b.h.,
garia. The two populations have sometimes been straight or curved. Bark on trunks of larger trees 733
viewed as two distinct varieties, with “Pinus peuce breaking into large, irregular, more or less smooth
var. vermiculata Christ” (cited in Vidaković, 1991: plates separated by broad and deep, blackish fis-
532 but orig. publ. not traced) in Bulgaria, but this sures. Branches long, spreading and ascending;
is not generally accepted as the stated differences: higher order branches in lower crown drooping,
length of leaves and cones, are continuous and part- forming a dense, rounded crown. Foliage branches
ly overlapping. Extensive exploitation in the past stout; new shoots green with brown-red spots, turn-
has undoubtedly had an impact on its abundance ing to pinkish brown on the upperside and fading
but no details of a possible reduction are known. At to grey-brown. Buds large; terminal bud 20–35 mm
present, this species occurs in several protected ar- long, 8–12 mm wide, oblong-conical, not resinous;
eas in the Balkan mountains. cataphylls red-brown, reflexed. Leaves in fascicles of
IUCN: NT 2, held by a persistent, 20–30 mm long, dark grey
basal sheath, remaining on branchlets 2–3 years,
straight or sometimes slightly curved, rigid, 10–20
Uses
(–25) cm long, not or slightly twisted, 1.5–2 mm
Balkan pine is a minor timber tree in the Balkans; it wide, lustrous grey-green or dark green; margins
is now commonly planted for forestry in the region denticulate; apex acute or pungent; stomata in con-
and natural stands are less intensively exploited than spicuous lines on all faces. Pollen cones cylindrical,
in the past. The wood is used only locally for carpen- 2–4 cm long, yellow. Seed cones solitary or more of-
try and furniture and can be rather knotty, but when ten in whorls of 2–4, short pedunculate, persistent
plantation-grown trees come of age to be harvested, several years, spreading or reflexed, narrowly ovoid-
this tendency will be reduced. Inter-specific hybrids conical when closed, 10–20(–22) cm long and 5–8
with other species in subsection Strobus have been cm wide when closed, not opening soon but usually
tested in forestry nurseries for growth performance. open before falling. Seed scales thick woody, rigid,
Balkan pine, in contrast with several other species, small and imbricate near the slightly oblique cone
is resistant to the pathogenic fungus Cronartium base, much larger and longer beyond, spreading
ribicola (Basidiomycota) and hybrids may also be- wide, oblong, more or less flat. Apophyses broadly
come resistant to the disease. In horticulture it has rhombic in outline, raised and thickened to pyrami-
long been used, being first introduced north of the dal, prominently transversely keeled and mostly
Alps in Germany in 1863. It is mostly grown as a also with 2–3 raised longitudinal ridges, lustrous
‘species tree’ for large gardens and parks; only a lim- orange-brown or red-brown; umbo dorsal, central,
ited number of cultivars, mostly producing dwarfed forming a straight or forward curved, short spine.
forms, is known. Seeds obovoid-­oblong, (7–)8–10(–12) mm long, grey
or blackish grey (sometimes different on two sides);
wing 22–30 mm long, 7–10 mm wide, oblique.
Pinus pinaster Aiton, Hort. Kew. 3: 367. 1789.

Etymology Taxonomic notes

Pinastre (also pinastro) is one of the vernacular This species is divided into three subspecies in several
names for this pine, which was adopted for the spe- recent European standard Floras. Their morpholog-
cies epithet by William Aiton. ical differences are small and are probably to some
 extent due to differences in climate and/or habitat. century. It has been planted to stabilize dunes in
These subspecies are listed below. There has long coastal areas within as well as far outside its natural
been a keen interest in forestry to discover differ- range where frost is a minimal risk due to the mari-
ences in growth, habit, and frost resistance between time influence on the microclimate. The region Les
the two provenances represented by P. ­pinaster sub- Landes in SW France is one of the principal areas
sp. pinaster (“var./subsp. atlantica”, an illegitimate where this pine has been planted for forestry purpos-
name) and subsp. escarena (P. meso­geensis of several es, including resin tapping. Another centre for resin
authors) from the Atlantic and Mediterranean coasts production is Corsica. It has also been planted on
respectively. This excludes the upland North African a forestry scale in many parts of the Mediterranean
taxon, P. pinaster subsp. renoui; these rare trees were region and in South Africa, where it has become one
734 never in the focus of foresters. It may be, on closer of the invasive species of pine as a result. The wood is
scrutiny, the only subspecies (or variety) worthy of of coarse grain and very resinous. It is (or was) used
taxonomic recognition, as it has more slender leaves for mining pit props, construction, telephone poles
with only 2 resin ducts and smaller seed cones. The and posts for fencing, boat-building, and furniture.
distinguishing morphological character states cited Smaller sized logs with many knots or blemishes
for the other two are, to the contrary, not discon- are usually chipped or pulped for particleboard and
tinuous and even largely overlapping; the taxonomic paper. The large volume of bark per unit of wood
case for their separation is therefore less convincing. makes this a pine suitable for production of mulch.
Resin is extracted by tapping as well as distillation
of bark and wood pulp, and used to produce tur-
Distribution
pentine, pitch, oils, varnishes, waxes and soap. The
SW and S Europe; N Africa, Morocco, border be- large, decorative cones are sold as ornamental pieces
tween Algeria and Tunisia. in flower displays and for Christmas decorations. In
TDWG codes: 12 BAL COR FRA-FR POR SAR SPA- horticulture it is less common and most suitable for
SP 13 ITA-IT SAR SIC-SI 20 ALG MOR-MO TUN dry, sandy soils; only a few cultivars are known.

3 subspecies are recognized:

Ecology
Pinus pinaster is mainly a pine of low-lying, coast- Pinus pinaster Aiton subsp. pinaster. Pinus pinaster
al plains, usually on sandy soils of sea shore dunes Aiton subsp. atlantica Villar, Bol. Soc. Esp. Hist.
and flats; however, in Morocco this species extends Nat. 33: 427. 1934. Type not designated. Fig. 240, 241
into the mountains to an elevation of ca. 2000 m.
Extensive planting, e.g. in sand dunes, for centuries Pinus pinaster Aiton var. acutisquama Boiss., Voy.
has established this species in areas where it may Bot. Espagne 2: 583. 1841; Pinus pinaster Aiton subsp.
not have occurred naturally but where it has sub- acutisquama (Boiss.) Rivas-Martínez, Rivasgodaya
sequently often been naturalized. It forms tall pine 6: 52. 1991.
forests and if with a more or less open canopy the
understorey is of evergreen maquis shrubs. This spe-
Description
cies is frost sensitive, which is probably a reason why
only in the far south of its range it ascends into the Trees to 30 m tall. Leaves 10–20 cm long; resin ducts
mountains. In Morocco it is a constituent of mixed at base of leaves (within sheath) limited to the two
coniferous forest with Pinus nigra subsp. salzmannii, lateral, larger ducts, at mid-leaf several additional
Abies ­pinsapo var. marocana, Cedrus atlantica, and small ducts; seed cones 10–18 cm long.
Taxus baccata. Common angiosperm trees in this
forest type are Quercus ilex and at the higher alti-
Distribution
tudes Populus tremula.
France (Atlantic coast), Portugal, Spain; natural
distribution uncertain due to long-time extensive
Uses
planting.
Maritime pine is an important timber tree as well TDWG codes: 12 FRA-FR POR SPA-SP
as Europe’s major source of turpentine since the 16th
 Conservation Ecology
IUCN: LC In Morocco this subspecies is reported to occur at
elevations up to 2000 m in the mountains, which is
Pinus pinaster Aiton subsp. escarena (Risso) unusual for the species.
K. Richt., Pl. Europ. 1: 1. 1890. Pinus escarena Risso,
Hist. Nat. Europ. Mérid. 2: 340, 459. 1826. Type not
Conservation
designated.
This subspecies of Pinus pinaster is very rare and
Pinus mesogeensis Fieschi & Gaussen, Bull. Soc. known from only two widely separated popula-
Hist. Nat. Toulouse 64: 440. 1932; Pinus pinaster Ai- tions on the African Mediterranean coast, the larg- 735
ton var. mesogeensis (Fieschi & Gaussen) Silba, Phy- est being in Morocco. These are relict occurrences
tologia 68: 59. 1990. and may indeed be genetically distinct. Logging and
habitat degradation are known to occur, but to what
extent is presently unknown. Afforestation with oth-
Description
er pines, especially with P. pinaster from European
Trees to 40 m tall. Leaves up to 25 cm long, 2 mm provenance, threatens the genetic integrity of these
wide; resin ducts at base of leaves (within sheath) relict populations.
usually more than the two larger lateral ducts, at IUCN: EN (B 2, a [iii])
mid-leaf numerous; seed cones up to 22 cm long.

Pinus pinceana Gordon, Pinetum: 204. 1858. Type:

Distribution
Mexico: Morelos, Cuernavaca, C. A. Ehrenberg s.n.
W Mediterranean: France, Spain, Baleares, Corsica, (holotype not located, isotype W). Pl. 32
Sardinia, Italy, Sicily, Malta.
TDWG codes: 12 BAL COR FRA-FR ITA-IT SAR SIC-
Etymology
MA SIC-SI SPA-SP
This species was named after “Mr Pince of Exeter”,
presumably a nurseryman.
Conservation
IUCN: LC
Vernacular names
Pinus pinaster Aiton subsp. renoui (Villar) Maire, Weeping pinyon pine, Pince’s pinyon pine; piñon
[Fl. Afrique N.] Encycl. Biol. 33: 145. 1952. Pinus (Spanish)
pinaster Aiton var. renoui Villar, Vol. Jubil. Soc.
Sci. Nat. Maroc: 241. 1948; Pinus renoui (Villar)
Description
Gaussen, Rev. Int. Bot. Appl. Agric. Trop. 32: 515.
1952. Type not designated. Trees or large shrubs to 6–10(–12) m tall, d.b.h. to 20–
30 cm. Trunk short, often branching from near the
ground, erect, monopodial, contorted or forked. Bark
Description
on lower part of trunk and lowest branches exfolia-
Trees to 25 m tall, crown umbrella-shaped or domed, ting into irregular plates, brownish grey. Branches as-
dense. Leaves 12–20 cm long, slender, 1.5–2 mm cending or spreading, of higher orders long, slender,
wide, with only 2 lateral, large resin ducts at mid- flexible, the ultimate branches drooping to pendu-
leaf. Seed cones 10–15 cm long. lous, forming a broad, irregular, open crown. Shoots
slender, flexible, glabrous and smooth, brownish grey
to grey. Cataphylls 3–4 mm long, light brown. Veg-
Distribution
etative buds ovoid-globose to ovoid-­conical, to 4 mm
N Africa: Morocco, and on Algerian-Tunisian bor- long, not resinous. Fascicle sheaths ca. 10 mm long,
der near the coast. soon spreading or curling and deciduous. Leaves in
TDWG codes: 20 ALG MOR-MO TUN fascicles of 3, rarely 4, ­remote, ­persisting 2–3 years,
736

1
3

2 5

Pl ate 32 . Pinus pinceana. 1. Habit of tree. 2. Branch with foliage and seed cone. 3. Leaves. 4. Seed cone.
5. Seed.
straight, rigid, 5–12(–14) cm long, 0.8–1.2 mm wide, and far apart and usually with a low crop of cones
with entire margins, acute, greyish green. Stomata compared to P. cembroides and its close relatives. It is
mostly in 2–4(–5) lines on each adaxial face. Pollen not known in cultivation outside a few botanic gar-
cones ovoid-­oblong, 8–10 × 4–5 mm, purplish or yel- dens and arboreta; it will be suitable in gardens with
lowish. Seed cones lateral, not on ultimate branches, a Mediterranean climate or nearly equivalent.
solitary or rarely in pairs on slender, curved and easily
breaking peduncles, ovoid-oblong to short cylindri-
cal, often irregular, 5–10 × 3.5–6(–7) cm when open. Pinus pinea L., Sp. Pl. 2: 1000. 1753. Type: Illustra-
Seed scales parting only slightly or sometimes more tion: “Pinus ossiculis duris, foliis longis” in Bauhin &
widely, easily moveable, thick woody, concavo-con- Cherler, Hist. Pl. Univ. 1 (2): 248. 1650 (lectotype).
vex, with (1–)2 deep, cup-like depressions holding Fig. 242, 243 737
the seeds. Apophysis irregular, prominently raised,
transversely keeled, rhombic to pentagonal in out-
Etymology
line, with angular or undulating upper margin, lus-
trous red-brown, sometimes with concentric colour The species epithet is a Latin adjective (with femi-
rings or radially striated. Umbo dorsal, flattened at nine gender) meaning ‘of pines’ (masc. = pineus).
apex, up to 5 mm high, with a minute prickle. Seeds
obovoid, 11–14 × 7–8 mm, wingless when detached
Vernacular names
from the scale, brown.
Stone pine, Umbrella pine; pino domestico, pino
da pinoli (Italian); pino piñonero (Spanish); pin-
Distribution
hero manso (Portuguese); pin pignon, pin parasol
Mexico: Coahuila, Hidalgo, Querétaro, San Luis Po- (French)
tosí, Zacatecas.
TDWG codes: 79 MXE-CO MXE-HI MXE-QU MXE-
Description
SL MXE-ZA
Trees to 20–25 m tall; trunk to 1 m d.b.h., mono­
podial or soon forked into more stems. Bark scaly,
Ecology
becoming thick on trunk, breaking into large, hard
This species occurs often on calcareous slopes and plates divided by deep, irregular, dark brown to black
in ravines (barrancas) at altitudes between 1400– fissures, orange-brown to reddish brown with grey
2300 m a.s.l. in arid and semi-arid mountains of NE patches of flakes that remained longer. Branches nu-
Mexico. Annual precipitation is lower than in the merous, spreading and ascending, the higher order
Pinyon-Juniper belt (ca. 300–400 mm according to branches assurgent to erect, forming an umbrella-
Perry, 1991) and is concentrated in the summer. This shaped, dense crown; young trees are more or less
species sometimes occurs with Pinus cembroides, conical with an erect stem. Foliage branches slen-
more often with Juniperus flaccida, and usually with der or stout, new shoots glabrous, rough with pul-
numerous sclerophyllous or deciduous shrubs, e.g. vini of fallen leaf fascicles, pale green to yellowish
Prosopis (in arroyos), Mimosa, Karwinskia, Leucae­ brown, turning orange-brown. Buds ovoid-conical,
na, Sophora, Quercus, Cercocarpus, Gochnatia, and 6–12 mm long, acute, not resinous; cataphylls light
Fouquieria, as well as the succulent genera Yucca, brown or red-brown, with fringed margins and re-
Agave and Opuntia. The trees are often widely scat- flexed apex. Leaves in fascicles of 2 held by a short,
tered in this vegetation. persistent basal sheath, remaining 3–4 years, rigid
and spreading, 8–12(–15) cm long, usually twisted,
Conservation 1.2–1.8 mm wide; margins minutely serrate; leaf col-
our on young plants glaucous, becoming lustrous
IUCN: NT
dark green; stomata in fine lines on all surfaces.
Pollen cones short cylindrical, yellow. Seed cones
Uses
solitary or sometimes in whorls of 2–3 on short pe-
This species is not commercially used; the seeds are duncles, persisting several years, serotinous or semi-
edible (‘piñones’), but trees are scarce, inaccessible serotinous, ripening in the third year, symmetrical,
 broadly ovoid to subglobose, (8–)9–13(–15) cm long, timber is poor, ­being coarse and resinous and sel-
8–10 cm wide when closed, when finally opening up dom straight for any substantial length. It is locally
to 13 cm wide near the depressed base, leaving some used to make furniture. The true economic value of
basal scales on branch when falling. Seed scales thick this pine has since ancient times mostly resided in
woody, rigid, with 2 deep depressions on adaxial the edible seeds. These are harvested while still in-
side holding the seeds. Apophysis (slightly) raised, side the closed cones, which are pulled down with
angular in outline, convex with 4–6 ridges converg- hooked poles; the cones open when heated. For this
ing on the dorsal, central, obtuse umbo, maturing purpose lower tree crowns are favoured, which is ac-
from olive green to lustrous orange-brown or red- complished by keeping the canopy open. Seeds are
brown, weathering grey. Seeds obovoid, 15–20(–22) a delicacy in themselves, but are also used in recip-
738 mm long, 8–10 mm wide, brown; wing mostly rudi- ies ranging from pasta dishes to pastries. Millions of
mentary, oblique, 3–10(–15) mm long. kilogrammes of seeds or ‘pine kernels’ are harvested
in the Mediterranean countries each year. Empty
pine cones are good, hot burning fuel (bakeries), but
Distribution
can also be sold as ornamental objects to florists. The
Mediterranean Europe and Near East (doubtfully resin is tapped and used for medicinal treatments,
native in many areas of E Mediterranean). wax for violin bows, varnishes, and waterproofing.
TDWG codes: 12 BAL COR FRA-FR FRA-MO POR A green dye is made from the needles. The species is
SAR SPA-SP 13 ALB GRC ITA-IT KRI SIC-SI 34 CYP also much valued as an ornamental tree in gardens
EAI LBS-LB TUR and parks around the Mediterranean and in other
parts of Europe with mild winters. In South Africa it
has become naturalized and is an invasive and nox-
Ecology
ious species in the fynbos biome of the Cape.
This is primarily a pine of coastal areas in the Medi-
terranean, at elevations from sea level to 600 m, on
coastal dunes and flats as well as on lower slopes of Pinus ponderosa Douglas ex C. Lawson, Agric.
mountains and in the hills. Many present-day stands Man.: 354. 1836.
are the result of historic plantings, some going back
to Roman times, and if managed well, these can have
Etymology
a natural understorey of maquis scrub or mixture
with smaller broad-leaved trees. Mature trees have a The species epithet (Latin, fem. of ponderosus)
thick, fire resistant bark and the massive cones take means ponderous or massive.
three years to mature and are serotinous or semi-
serotinous. Seeds are nearly wingless and dispersed
Vernacular names
by birds (also eaten by rodents) or may scatter after
fire burned off the undergrowth and its heat assist- Ponderosa pine, Western Yellow pine
ed in opening the cones. Pinus pinea is usually an
emergent tree above shrubs (maquis) or in low, open
Description
forests; it can also occur with Pinus halepensis and in
Quercus ilex maquis-woodland. Trees to 65(–72) m tall, but in substantial parts of
its range 20–30 m is more commonly the maxi-
mum height; trunk to 2.5 m d.b.h., usually forming
Conservation
a straight, columnar bole. Bark on trunk breaking
IUCN: LC up into large to very large (30–40 cm diam.) ir-
regular or oblong plates divided by broad fissures,
exfoliating in small chips, bright orange-yellow or
Uses
reddish brown with darker brown or blackish fis-
Although depending on stand density the Stone sures. Branches persisting or self-pruning, spread-
pine may develop a tall, straight bole, most com- ing and assurgent, often long, with lower branches
monly its branches spread out and the trunk re- down-curved, contorted or more or less pendulous,
mains short. The quality of the wood for sawn forming broad, columnar or domed, open crowns.
Foliage branches stout; new shoots 0.7–2 cm thick, (Lauria, 1996). Only P. ponderosa var. scopulorum
rough with pulvini from fallen leaf fascicles, orange- is here recognized as a botanical variety within the
brown, becoming brown. Buds ovoid, to 2 cm long; species; P. arizonica is treated as a distinct species
terminal bud to 1 cm wide, all resinous; cataphylls largely, but not exclusively, occuring in Mexico. If we
red-brown with white fringes. Leaves in fascicles of include the smaller trees of the ‘North Plateau Race’
2–3, persisting 2–3 years, held by a ca. 15–20(–25) in P. ponderosa var. ponderosa (as we must), as well
mm long, persistent basal sheath, spreading, straight as the spurious taxon P. washoensis and the tall trees
or curved, 15–25 cm long, rigid or pliant, slightly of the Cascades – Sierra Nevada, then the only mor-
twisted, (1–)1.2–1.8 mm wide; margins minutely ser- phological difference that appears to hold up more
rulate; apex acute or acuminate; leaf colour green; or less consistently is the number of leaves per fasci-
fine lines of stomata on all surfaces. Pollen cones cle: predominantly 2, rarely 3 in var. ponderosa, 2–3 739
spirally arranged, erect and standing out radially, (with a N-S cline towards increasing frequency of
3–4.5 cm long when full-grown, ellipsoid-cylindric, 3) in var. scopulorum. This is a difference that barely
initially bright red or magenta, turning yellow at an- merits taxonomic recognition in conifers.
thesis from yellow pollen sacs. Seed cones solitary
or rarely in pairs on short peduncles to nearly ses-
Distribution
sile, leaving some scales on branch when falling,
nearly symmetrical, ovoid-conical when closed, W North America, from S British Columbia to just
ovoid to subglobose when opened and 5–12 cm long. south of the Mexico-USA border.
Seed scales oblong, spreading wide or reflexed, thin TDWG codes: 71 BRC 73 COL IDA MNT ORE WAS
woody, rigid. Apophyses slightly to prominently WYO 74 NDA NEB OKL SDA 76 ARI CAL NEV UTA 77
raised, transversely keeled, more or less rhombic in NWM TEX 79 MXE-CO MXN-SO
outline, dull tan or lustrous reddish brown; umbo
raised (rarely flat), pyramidal, keeled, recurved and
Ecology
usually armed with a hooked prickle. Seeds ellipsoid
or obovoid, (4–)5–7(–8) mm long, brown or light Pinus ponderosa is one of the most widely distributed
brown, often mottled darker; wing 15–25 mm long. pines, with a range spanning 20 degrees of latitude
and from the lowlands to 3300 m a.s.l. While in the
north and east its limites are defined by temperature
Taxonomic notes
and rainfall respectively, in the south it is replaced
Pinus ponderosa is morphologically variable mainly by related species. The lower limit of average annual
in a north-south gradient from Canada to Mexico, precipitation is around 300 mm, the upper limit 1750
but also from the Pacific coast to the Rocky Moun- mm. It grows on soils derived from many rock types,
tains. Its economic importance has stimulated nu- both acidic and basic varying from around pH 5 to
merous studies into this variation, but much of it has pH 9, and usually with a capacity to retain mois-
not been built on a sound taxonomic basis. It has ture but well drained. Its great altitudinal range as
appeared from taxonomic studies, that what David well as geographic spread causes huge variations in
Douglas collected as P. ponderosa is called by for- temperature, both summer and winter, but to thrive
esters the ‘North Plateau Race’ of that species and and compete well it requires ample sunshine in the
is conspecific with P. washoensis, described from growing season. Pinus ponderosa is the first tall pine
Lake Tahoe in Nevada. The Ponderosa pine of the to appear above Pinyon-Juniper woodland in the in-
Cascade Range in Oregon and the western slopes terior of the western USA, but it is a seral species in
of the Sierra Nevada in California, while here in- the mesic mixed conifer forests of the Cascades and
cluded in P. ponderosa (‘Pacific Race’ of foresters), Sierra Nevada, where other conifers will eventually
would, if sufficiently different and distinct as a spe- dominate. It grows with numerous other conifers
cies from P. washoensis, have to be called P. bentha­ (for listings see under P. monticola and P. jeffreyi) in
miana. American foresters treated this pine as if it these mountain ranges. Its relatively high tolerance
included the type of P. ponderosa. Nobody seems of fire assures it a place in the succession in natural
to have bothered to ask what the type specimen of forests, but selective logging as well as fire preven-
P. ponderosa might be and where it had been col- tion have altered the forest composition to the dis-
lected; these questions have now been resolved advantage of P. ponderosa in many managed forests.
 Uses Conservation
Ponderosa pine is one of the most important tim- IUCN: LC
ber trees in the USA. Its wood properties and po-
tential size make it ideally suitable for sawn timber Pinus ponderosa Douglas ex C. Lawson var. scopu­
used in construction. It is used as roundwood for lorum Engelm., in Watson, Bot. California 2: 126.
posts and sawn for crates as well as made into par- 1880. Pinus scopulorum (Engelm.) Lemmon, Gard.
ticle board and pulp. Timber of large sizes and high & Forest 10: 183. 1897; Pinus ponderosa Douglas ex
grade is sawn for light construction like window C. Lawson subsp. scopulorum (Engelm.) E. Murray,
frames, doors, stairs, flooring, sidings, panelling, Kalmia 12: 23. 1982. Type: USA: Colorado [“Colo-
740 veneers and for furniture, cabinetwork, boxes, and rado Territory, lat. 39–41°, Alpine and Subalpine”],
woodware. In the National Forests of the USA the C. C. Parry s.n. (lectotype MO).
recreational and ecological functions of the open,
sunny forests of Ponderosa pine at relatively low
Description
altitudes are increasingly recognized and on public
lands these trees are no longer exclusively regarded Leaves in fascicles of 2–3. Seed cones 5–10 cm long;
as a source of timber. The species and its variety seeds 3–5 mm long; seed wings 12–15 mm long (seed
scopulorum are common in horticulture in parts of character states are correlated with cone size and
Europe, but usually limited to large parks and arbo- smaller in this variety because the cones are ­smaller).
reta. A few cultivars have been named, but they are
uncommon. Introductions for forestry plantations
Distribution
have been less succesful due to fungal diseases, e.g.
in New Zealand. USA: Rocky Mts., Black Hills; Mexico: near the bor-
der in Coahuila and Sonora.
2 varieties are recognized: TDWG codes: 73 COL MNT WYO 74 NDA NEB OKL
SDA 76 ARI NEV UTA 77 NWM TEX 79 MXE-CO
Pinus ponderosa Douglas ex C. Lawson var. pon­ MXN-SO
derosa. Type: ovuliferous cone with appended label
stating: “Bayford – Bury January 2° – 1849” (neo-
Conservation
type W). Fig. 244
IUCN: LC
Pinus washoensis Mason & Stockw., Madroño 8: 62.
1945; Pinus ponderosa Douglas ex C. Lawson subsp.
washoensis (Mason & Stockw.) E. Murray, Kalmia 12: Pinus praetermissa Styles & McVaugh, Contr.
23. 1982. Univ. Michigan Herb. 17: 310, f. 2. 1990. Type:
Mexico: Nayarit, Jalá, Cerro Juanacata, J. W. Stead
& B. T. Styles 475 (holotype FHO).
Description
Leaves mostly in fascicles of 2, rarely of 3. Seed cones
Etymology
8–12 cm long; seeds 6–8 mm long; seed wings 15–25
mm long (seed character states are correlated with The species epithet means ‘overlooked’ or ‘neglected’
cone size, not independent character states, and and refers to the fact that this species was previously
larger in this variety because the cones are larger). not recognized as distinct.

Distribution Vernacular names

Pacific W North America, from S British Columbia No vernacular names have been recorded for this
to S California. species.
TDWG codes: 71 BRC 73 IDA MNT ORE WAS 76 CAL
NEV
 investigation is necessary to evaluate its sympatry
Description
with these and other pines.
Trees to 10–15(–20) m tall, d.b.h. to 30 cm; trunk mo-
nopodial, tortuous or curved, sometimes branching
Conservation
low. Bark irregularly fissured longitudinally, red-
dish brown. Branches long, spreading, tortuous, not IUCN: LC
pendulous, forming a broad, irregular, open crown.
Shoots slender, reddish brown, turning greyish.
Uses
Cataphylls 7–10 mm long, subulate, not recurved,
scarious, light brown. Vegetative buds oval-oblong This scattered species is rarely recognized as distinct
to cylindrical; terminal bud 10–15 mm long; lateral from P. oocarpa by loggers and consequently no spe- 741
buds ovoid-acute, smaller, not resinous. Fascicle cific uses are known. It is likely to be exploited for
sheaths 10–14 mm long, persistent, weathering dark timber together with other species. It is a subtropical
brown to grey. Leaves in fascicles of (4–)5, persist- species not known in horticulture.
ing up to 3 years, straight, delicate and lax but not
pendulous, (8–)10–16 cm long, 0.5–0.8 mm wide,
serrulate at margins, acute, bright green. Stomata on Pinus pringlei Shaw, in Sargent, Trees & Shrubs
all faces of the leaves. Pollen cones ovoid-oblong to 1: 211, t. 100. 1905. Type: Mexico: Michoacan,
cylindrical, 1–1.5 cm × 5 mm, pink to reddish. Seed Uruapán, Cerro de la Cruz, [“hilltops near
cones subterminal, commonly solitary, occasionally Uruapán”], C. G. Pringle 10019 (lectotype GH).
opposite, on up to 35 mm long, recurved, slender
peduncles, deciduous, non-serotinous, falling with-
Etymology
out basal scales and peduncle in most cases. Ma-
ture cones broadly ovoid to subglobose (wider than This species was named after Cyrus G. Pringle
long), often with a flattened base when opened and (1838–1911), who made many collections of plant
missing the proximal scales, (4–)5–6.5(–7) × (5–)6– specimens in Mexico.
8 cm when open. Seed scales thin woody but rigid,
oblong, straight or recurved, more or less symmetri-
Vernacular names
cal. Apophysis flat to slightly raised, radially striate
or ridged to transversely keeled, rhombic to pen- Pringle’s pine
tagonal in outline, lustrous light brown or yellowish
brown. Umbo dorsal, flat or slightly raised, obtuse.
Description
Seeds obliquely ovoid, 5–8 × 3–4 mm, blackish grey
or with black dots; wings 12–18 × 5–8 mm, greyish Trees to 20–25 m tall, d.b.h. to 1 m; trunk monopo-
brown. dial, straight. Bark thick, rough, with small, scaly
plates, fissured; inner bark orange-red; outer bark
greyish brown. Branches spreading or arching, assur-
Distribution
gent near the top, forming an open, broadly domed
Mexico: Durango, Jalisco, Nayarit, Sinaloa. or irregular crown. Shoots thick, smooth, reddish
TDWG codes: 79 MXE-DU MXN-SI MXS-JA MXS- brown, glaucous. Cataphylls lanceolate to subulate,
NA 10–12 mm long, scarious, becoming reflexed at apex,
reddish brown. Vegetative buds ovoid-oblong to cy-
lindrical; terminal bud 10–15 mm long; lateral buds
Ecology
ovoid-acute, smaller, not resinous. Fascicle sheaths
Pinus praetermissa occurs in open, dry pine-oak up to 20 mm long, persistent but reduced to 10–15
woodlands or tropical broad-leaved forests, often mm length, dark red-brown to blackish. Leaves in
on rocky slopes. Its altitudinal range is 900–1900 m fascicles of 3(–4), persisting 2–3 years, rigid, straight,
a.s.l. The dry season is from November to May, an- (15–)18–25(–30) cm long, 1–1.5(–1.7) mm wide, with
nual precipitation varies between 1000–1500 mm. serrulate margins, acute-pungent, lustrous (yellow-
Associated pines are P. devoniana, P. lumholtzii, ish) green or glaucous green. Stomata on all faces of
P. pseudostrobus, and possibly P. oocarpa, but further the leaves. Pollen cones ovoid-oblong to cylindrical,
 1.5–2.5 cm × 8 mm when shedding pollen, yellowish. Pinus pseudostrobus Lindl., Edwards’s Bot. Reg. 25:
Seed cones subterminal, solitary or more commonly 63. Aug 1839. [Allg. Gartenzeitung 7: 325. 1839]. Pl. 33
in whorls of 2–4 on short, stout, curved and tena-
cious peduncles. Mature cones ovoid, slightly curved
Etymology
or asymmetrical, 5–8(–10) × 3.5–6(–7) cm when
open, but semi-serotinous (only partially opening), The species epithet refers to its (superficial) similar-
leaving some basal scales on branch when falling. ity with P. strobus.
Seed scales thick woody, oblong, almost straight,
symmetrical. Apophyses on one side of cone nearly
Vernacular names
flat to slightly raised, on other (outer) side slightly
742 more pronounced, these transversely keeled, rhom- Smooth-bark Mexican pine, False Weymouth pine;
bic to pentagonal in outline, lustrous ochraceous or pino blanco, pino chalmaite, pino lacio, pino liso
light brown. Umbo dorsal, flat or depressed, with a (Spanish)
minute, deciduous prickle. Seeds obovoid, 4–6 mm
long, dark brown to grey-black; wings 14–18 × 6–8
Description
mm, translucent, light brown.
Trees to 40(–45) m tall, d.b.h. to 1 m; trunk mo-
nopodial, erect. Bark thick, scaly, with elongated
Distribution
plates and deep, longitudinal fissures, dark brown to
Mexico: Michoacán, México, Morelos, Guerrero grey-brown. Branches spreading or ascending; low-
and Oaxaca, perhaps in W Puebla. er branches curved down; upper branches ascend-
TDWG codes: 79 MXC-ME MXC-MO MXS-GR MXS- ing. Shoots slender, smooth, glaucous or pruinose.
MI MXS-OA Cataphylls 10–15 mm long, subulate-caudate, soon
recurved, reddish brown to dark brown. Vegeta-
tive buds ovoid-conical, acute; terminal bud 15–20
Ecology
× 10–15 mm; lateral buds smaller, not resinous. Fas-
This is a pine of the montane, mesic forests of the cicle sheaths (15–)20–30(–35) mm long, persistent,
Sierra Madre del Sur and parts of the ‘Eje Volcáni- lustrous red-brown, weathering grey-brown. Leaves
co Transversal’, where it is growing between 1500– in fascicles of 5, rarely 4 or 6, persisting 2–3 years,
2600(–2800) m a.s.l. This region has a subtropical slender, straight, spreading or drooping, usually lax,
to warm temperate climate; annual precipitation (18–)20–30(–35) cm long, 0.8–1.3 mm wide, with
varies between 1000–2000 mm, mainly occurring serrulate margins, acute, (light) green or glaucous
in summer storms from June through September. It green. Stomata on all faces of leaves. Pollen cones
is a constituent of pine and pine-oak forests; associ- ovoid-oblong to cylindrical, 20–35 × 5–7 mm, pur-
ated pines include Pinus douglasiana, P. maximinoi, plish or pinkish yellow. Seed cones subterminal,
P. pseudostrobus, P. oocarpa, and P. patula. On drier solitary or in pairs, more rarely in whorls of 3–4 on
sites, often in degraded or secondary forest, P. law­ very short, stout peduncles remaining with a few
sonii and P. devoniana are more commonly growing cone scales on branch when cone has fallen. Mature
with P. pringlei; in both forest types Quercus spp. are cones more or less asymmetrical, often curved at
often (co-)dominant. Dvorak & Donahue (1992) re- base, ovoid-oblong to broadly ovoid when opened,
port a grass stage of the seedlings of P. pringlei. then 7–16 × 6–13 cm. Seed scales usually thick
woody, oblong, straight or slightly curved. Apo­
physes extremely variable, from nearly flat to elon-
Conservation
gated, more so on one side of cone and towards base,
IUCN: LC transversely keeled, tapering to an obtuse umbo (or
umbo mucronate), rhombic or pentagonal in out-
line, upper margin angular, irregularly undulate, or
Uses
rounded; colour in various hues of brown. Umbo
This species is reported to have dense wood and is dorsal, variable, from obtuse to prominent, 3–15 mm
used for lumber; the resin is also tapped commer- long, without a prickle or prickle deciduous. Seeds
cially. It is not known to be in horticultural use. 5–7 × 3–4.5 mm, ochraceous to grey-brown, with
 5
743

2
1

4
3

Pl ate 33. Pinus pseudostrobus. 1. Habit of tree. 2. Branchlets with foliage. 3. Leaves. 4. Seed cone (var.
pseudostrobus). 5. Seed cone (var. apulcensis).
 or without dark spots; wings obliquely-ovate, with has led to regional depletion of the best stands. The
a straight side, 20–25 × 7–10 mm, yellowish brown. wood is used for light construction, carpentry and
joinery, wall panelling, veneers, boxes and contain-
ers, matches, and wood pulp. In Mexico, this species
Distribution
is also used as a source of resin. This tree has been
Mexico: Sinaloa-Durango border, Nuevo León, SE widely planted for forestry in Africa and to a lesser
Coahuila, E Guanajuato(?), Jalisco, Michoacán, extent in India; in horticulture its is rare and gener-
México, Distrito Federal, Morelos, Hidalgo, Puebla, ally only suitable in regions with a mild climate.
Tlaxcala, W Central Veracruz, Guerrero, Oaxaca,
Chiapas; Guatemala (highlands); W Honduras; N El 2 varieties are recognized:
744 Salvador.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC- Pinus pseudostrobus Lindl. var. pseudostrobus.
PU MXC-TL MXE-AG MXE-CO MXE-CU MXE-DU Type: Mexico: Michoacan, Angangueo, C. T. Hart­
MXE-GU MXE-HI MXE-NL MXE-QU MXE-SL MXE- weg s.n. (lectotype P).
TA MXE-ZA MXG-VC MXN--SI MXN-SO MXS-CL
MXS-GR MXS-JA MXS-MI MXS-NA MXS-OA MXT-CI Pinus pseudostrobus Lindl. var. estevezii Martínez,
80 ELS GUA HON Anales Inst. Biol. Univ. Nac. México 16: 188. 1945; Pi­
nus estevezii (Martínez) J. P. Perry, J. Arnold Arbor.
63: 187. 1982.
Ecology
Pinus pseudostrobus Lindl. var. apulcensis (Lindl.)
This is a widely distributed and in many places com- Martínez, Anales Inst. Biol. Univ. Nac. México 16:
mon or abundant pine of montane to high montane 192. 1945, non Shaw (1909); Pinus pseudostrobus
habitat in cold-temperate to warm-temperate zones. Lindl. subsp. apulcensis (Lindl.) Stead, J. Linn. Soc.,
Its altitudinal range is considerable, but differs little Bot. 89: 269. 1984, excl. typus.
along the entire 2500 km of its range: (850–)1900– Pinus nubicola J. P. Perry, J. Arnold Arbor. 68: 447.
3000(–3250) m a.s.l. Annual precipitation is also 1987.
variable, but with a minimum of ca. 800 mm, in Pinus pseudostrobus Lindl. var. laubenfelsii Silba,
Guatemala and Honduras it can be over 2000 mm. Phytologia 68: 60. 1990.
Throughout its range it is an important constitu- Pinus yecorensis Debreczy & Rácz, Phytologia 78 (3):
ent of mixed conifer pine and pine-oak forests, oc- 15. 1995.
casionally associated with Liquidambar, as on wet Pinus yecorensis Debreczy & Rácz var. sinaloensis
E slopes on the Gulf coast side of the mountains of Debreczy & Rácz, Phytologia 78 (3): 18. 1995.
Central and S Mexico. In the many disturbed forests
it may survive as scattered groves or as individual
Description
trees, often with an understorey of e.g. Gaultheria,
Cassia or, when associated with fire, grasses and/or Apophysis variable, from nearly flat to prominently
Pteridium aquilinum. In its driest habitat in Central raised, more so on one side of cone and towards
and NE Mexico it occurs with P. cembroides, Juni­ base, transversely keeled, tapering to an obtuse
perus flaccida, and Quercus and an understorey with umbo, rhombic or pentagonal in outline; upper
e.g. Agave, Buddleja, Opuntia, and Salvia, probably margin angular, irregularly undulate, or rounded;
mostly in secondary forest. colour in various hues of brown. Umbo dorsal, ob-
tuse, 3–6 mm long, 4–6 mm wide at base, without a
prickle or prickle deciduous, usually darker than the
Uses
apophysis.
Pinus pseudostrobus is one of the most common
and important ‘hard pines’ in the southern half of
Distribution
Mexico and the highlands of Guatemala and parts
of Honduras. Its wood is light yellowish in colour, Mexico; Guatemala; Honduras; N El Salvador.
light weight (specific gravity 0.45), strong, with long TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC-
intervals of knot-free wood which is only slightly PU MXC-TL MXE-CO MXE-DU MXE-HI MXE-NL
resinous. Exploitation for timber is widespread and
MXG-VC MXN-SI MXS-GR MXS-JA MXS-MI MXS-OA Pinus pumila (Pall.) Regel, in Kuester & al., Index
MXT-CI 80 ELS GUA HON Sem. Hort. Bot. Imp. Petrop. 1858: 23. 1859. Pinus
cembra L. var. pumila Pall., Fl. Ross. 1 (1): 5, t. 2, f.
F-I. 1784. Type: Illustration in Pallas, Fl. Ross. 1 (1):
Conservation
t. 2, f. F-I. 1784 (lectotype, designated here).
IUCN: LC
Etymology
Pinus pseudostrobus Lindl. var. apulcensis (Lindl.)
Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 19, t. 12, The species epithet means ‘dwarfish’ or ‘low’ grow-
f. 6–8. 1909. Pinus apulcensis Lindl., Edwards’s Bot. ing and refers to its habit.
Reg. 25: 63. Aug 1839. [Allg. Gartenzeitung 7: 325. 745
1839]. Type: Mexico: Hidalgo, Apulco, C. T. Hartweg
Vernacular names
s.n. (lectotype W (cone), isolectotype GH).
Dwarf Siberian pine; Kiedrovnik (Russian)
Pinus oaxacana Mirov, Madroño 14: 145. 1958; Pi­
nus pseudostrobus Lindl. var. oaxacana (Mirov)
Description
S. G. Harrison, Taxon 14 (7): 247. 1965.
Pinus oaxacana Mirov var. diversiformis Debreczy & Prostrate or spreading, sometimes erect shrubs
Rácz, Phytologia 78 (3): 19. 1995. 1–4(–6) m tall, with layering branches and often
forming extensive thickets; rooted new stems as-
cending or assurgent. Bark thin, smooth, flaking
Description
on larger stems, grey-brown. Foliage branches slen-
Apophysis variable, prominently raised and partly der, flexible, not breaking easily, new shoots at first
elongated, especially on one side of cone and towards densely pubescent, later glabrous, initially grey-
base, transversely keeled, rhombic or pentagonal in green or brown, turning darker red-brown in the
outline; upper margin angular, irregularly undulate, second or third year, weathering grey-brown. Buds
or rounded; colour in various hues of brown. Umbo ovoid-conical, to 10 mm long, acute, resinous; cata-
dorsal, prominent and/or elongated, 5–15 mm long, phylls reddish brown. Leaves crowded in dense tufts
5–10 mm wide at base, without a prickle, usually at end of shoots, persisting 3–4 years, in fascicles of
darker than the apophysis. 5 with a deciduous basal sheath, 4–7(–8) cm long,
more or less rigid, spreading but curved forward,
ca. 1 mm wide, trapezoid in cross-section; margins
Taxonomic notes
minutely serrulate; leaf colour green abaxially, glau-
For a taxonomic assessment and the correct applica- cous green adaxially; stomata on the two adaxial
tion of this infraspecific name and its synonyms, see faces in white lines. Pollen cones crowded, spirally
Farjon (1995) and Farjon & Styles (1997). arranged, ovoid-ellipsoid, conspicuously red-purple
before anthesis. Seed cones solitary or in whorls of
2–4, short pedunculate or sessile, semi-persistent,
Distribution
remaining closed or the scales parting only slightly,
Mexico: México, Hidalgo, Puebla, Tlaxcala, W Cen- 3–5 cm long, ovoid to barrel-shaped, 2.5–3 cm wide,
tral Veracruz, Guerrero, Oaxaca and Chiapas; also green turning brown, resinous. Seed scales imbri-
in the Guatemala highlands and in N El Salvador. cate, rhombic to obovate, with 2 deep seed cavities
TDWG codes: 79 MXC-ME MXC-PU MXC-TL MXE- adaxially; apophyses broadly triangular, thickened,
HI MXG-VC MXS-GR MXS-OA MXT-CI 80 ELS GUA longitudinally striated or grooved, distal margins
slightly upturned; umbo (sub)terminal, distinct,
dark purple turning black. Seeds relatively large,
Conservation
7–10 mm long, obovoid, 5–7 mm wide, somewhat
IUCN: LC angular or with a ridged abaxial margin, dark brown
and wingless, remaining inside cone.
 Distribution Vernacular names
N Mongolia; Russian Federation: E Siberia, Russian Table Mountain pine, Prickly pine, Hickory pine
Far East (including islands); China: Inner Mongolia,
Manchuria (scattered); North & South Korea; Japan:
Description
Hokkaido, N Honshu.
TDWG codes: 30 BRY CTA IRK YAK 31 36 CHI-NM Small trees to 12–15 m tall; trunk to 60 cm d.b.h.,
CHM 37 MON 38 JAP-HK JAP-HN KOR-NK KOR-SK straight or crooked. Bark rough and scaly, breaking
into irregular plates, reddish brown or grey-brown.
Branches sparse, spreading widely, persistent, usu-
Ecology
746 ally forming a wide, flat-topped crown. Foliage
This is a pine well adapted to the extreme climate branches spreading and assurging, slender, rough
which prevails above the line of forests of Pinus syl­ with pulvini from fallen leaf fascicles, glabrous, yel-
vestris in the southern part of its range, while it re- lowish green turning yellowish brown to darker red-
places Larix gmelinii or birch forests at high altitude dish brown. Buds ovoid to short cylindric, up to 10
in the northern regions. It can be found scattered mm long, resinous, red-brown. Leaves in fascicles
in the understorey of these forests, too. Especially of 2, occasionally 3, held by a short, persistant ba-
on exposed mountain slopes close to the summer sal sheath, spreading, persisting 3 years, (3–)5–7(–8)
snowline it forms extensive, dense thickets. In Japan cm long, straight or more or less contorted, rigid,
the Dwarf Siberian pine occurs from 1400 m to 3200 1–1.5 mm wide, often twisted, green; margins serru-
m a.s.l., but on the Kamchatka Peninsula it is found late; apex acute to short acuminate; stomata in fine
from sea level up to 1200 m in favorable localities. Its lines on all surfaces. Pollen cones in dense clusters,
seeds are distributed by birds in the family Corvidae. spirally arranged, ca, 1.5 cm long, conspicuously red
when growing, turning yellow with the swelling of
pollen sacs and ripening of pollen. Seed cones lat-
Conservation
eral on branches, rarely solitary, usually in whorls
IUCN: LC of 3–6, sometimes more, nearly sessile or on short
peduncles, long persistent, variously serotinous but
eventually opening, asymmetrically ovoid-conical,
Uses
5–10 cm long, 4–6 cm wide, opening widest near
Dwarf Siberian pine is of little economic value. Its base to 8 cm. Seed scales thick woody, cuneate from
wood is of small, contorted size and shape and only base; apophyses prominently raised, diamond-
good for firewood in a region where there is plenty shaped at mid-cone, conical and recurved near base,
of this commodity of better quality. The seeds are ed- transversely keeled, largest on sun-exposed side of
ible, but difficult to harvest and mostly left to birds, cone, orange-brown or yellowish brown; umbo dor-
rodents and bears. In horticulture it is rarely met sal and nearly central, stout and armed with a dark,
with and mostly confined to botanic gardens and sharp spine. Seeds obovoid, somewhat angular and
arboreta, although in Russia and northern Japan it flattened, 5–6 mm long, blackish brown; wing 15–25
is also planted in some parks, road reservations and (–30) mm long, brown.
other amenity spaces. It should be a good species for
rock gardens in countries with cold winters. Distribution
E USA: Appalachian Mts.
TDWG codes: 75 PEN WVA 78 GEO MRY NCA SCA
Pinus pungens Lamb., Ann. Bot. (König & Sims) 2:
TEN VRG
198. 1805. Type not designated. Fig. 245
Ecology
Etymology
Pinus pungens is found in the Appalachian Moun-
The species epithet means ‘prickly’ and refers to the tains from the foothills to the crests, it becomes the
sharp spines on the umbos of the cone scales. dominant or single species of pine above 1300 m
a.s.l. in the southern part of its range. It grows on a
variety of soils, but is absent from limestone derived
Etymology
substrates. In the surrounding broad-leaved forests
it is restricted to outcrops or ridges of exposed rock. The species epithet refers to the common number of
Precipitation levels vary from north to south and four leaves in a fascicle.
with altitude between 760 mm and 2100 mm annu-
ally; summers are cool to warm while winters are Vernacular names
moderately cold and wet. This pine is a pioneer spe-
Parry pinyon, Nut pine; piñon (Spanish)
cies, able to take advantage of sudden environmen-
tal changes, e.g. disturbance by forest fire; it invades
Description
abandoned agricultural fields in many areas. In well
established stands in the mountains it is often mixed Trees or large shrubs to 10–15 m tall, d.b.h. to 30–50 747
with Pinus rigida or P. virginiana (at lower altitudes) cm; trunk short, erect, low branched or with a short,
and the angiosperms Acer rubrum, Nyssa sylvatica, clear bole. Bark thick, rough and scaly, with deep lat-
Quercus prinus, Q. coccinea, Q. velutina, Castanea eral and longitudinal fissures, reddish brown; outer
dentata, and Oxydendrum arboreum. Several species bark weathering grey. Branches spreading or ascend-
of Rhododendron, Vaccinium, Gaylussacia, and Kal­ ing, the higher order branches slender, spreading or
mia latifolia, Ilex montana and Smilax glauca often assurgent; crown rounded, becoming wide and open
form a dense understorey in stands of Pinus pungens in old trees. Shoots stout, rough with short decur-
on steep mountain slopes. rent pulvini and small cataphylls. Vegetative buds
ovoid-conical; terminal bud 6–10 × 4–5 mm; lateral
Conservation buds acute, orange-brown, slightly resinous. Fascicle
sheaths initially 5–8 mm long, soon breaking up in
IUCN: LC
recurving but early falling scales (not or rarely form-
ing rosettes). Leaves in fascicles of (3–)4(–5), rarely
Uses
a few fascicles 2 or 6; fascicles persisting (3–)4–7
Table mountain pine is of low value as a timber tree years; leaves curved or sometimes straight, rigid, of-
due to small size and ‘poor’ form, with crooked stems ten connivent for some time, (1.5–)2–4(–5) cm long,
and long branches. Pulpwood and firewood are the (0.8–)1–1.5(–1.7) mm wide, entire, acute-pungent,
only uses commonly cited to have any commercial dull or lustrous grey-green to glaucous green, with
value. Its worth to society is more of an ecological whitish stomatal bands. Stomata in two bands on
kind, as protective forest cover on unstable, shallow adaxial faces. Pollen cones ovoid-globose to short
rocky soils on mountain slopes and as habitat for cylindrical, ca. 10 mm long, purplish red, turning
wildlife. This species is not often seen in cultivation yellowish. Seed cones solitary or in whorls of 2–4 on
as it does not make a shapely tree; its gnarled form short, slender peduncles falling with cones. Mature
on rocky outcrops in nature can be very picturesque cones ovoid-globose to globose when closed, irregu-
and reminiscent of Chinese or Japanese landscape lar when opened, 4–6 × 4.5–7 cm when open. Seed
paintings. It is restricted to botanic gardens and ar- scales parting widely, moveable, irregular, concavo-
boreta in Europe and North America. convex, with 1–2 deep seed. Apophysis thick woody,
raised, pyramidal or obtuse conical, transversely
keeled, recurved or straight, ochraceous to yellow-
Pinus quadrifolia Parl. ex Sudw., USDA. Div. For- ish brown or reddish brown, often resinous. Umbo
est. Bull. 14: 17. 1897. Pinus cembroides Zucc. var. dorsal, flat or obtuse-pyramidal, centrally indented.
quadri­folia (Parl. ex Sudw.) Silba, J. Int. Conifer Pre- Seeds obliquely obovoid or elliptic, 12–18 × 8–12
serv. Soc. 7 (1): 29. 2000. Type: USA: California, San mm; integument thin (0.3–0.5 mm); megagameto-
Diego Co., “mountains E of San Diego” (Boundary phyte (‘endosperm’) white. Seed wings rudimentary
Survey under direction of major W. H. Emory), on the scale, absent from the developed, free seed.
C. C. Parry 1390 (holotype US). Fig. 246
Distribution
Pinus juarezensis Lanner, Southw. Naturalist 19 (1):
77. 1974; Pinus cembroides Zucc. var. juarezensis USA: California (Riverside and San Diego Co.);
(Lanner) Silba, Phytologia 68: 48. 1990. Mexico: Baja California Norte.
 TDWG codes: 76 CAL 79 MXN-BC
Description
Trees to 20–25(–33) m tall, d.b.h. to 2–2.2 m; trunk
Ecology
monopodial, often branched or forked near the
The altitudinal range of Pinus quadrifolia is 900– ground. Bark thick, rough and scaly, deeply fissured
2400(–2700) m a.s.l. It grows between the semi- vertically and horizontally, dark brown, weather-
desert and the chaparral scrub zones (and partly ing blackish grey. Branches spreading or ascend-
within the latter) and the mixed coniferous forest on ing, forming a dense, flat-topped or bushy crown.
the highest parts of the mountains. It is more widely Shoots multi-nodal, rough with prominent, decur-
distributed and often more common than P. mono­ rent pulvini and 5–10 mm long cataphylls. Vegeta-
748 phylla in the Pinyon-Juniper woodland, but occurs tive buds ovoid-acute; terminal bud 10–15 mm long;
often with it. Pinus jeffreyi is the only other pine with lateral buds smaller, all not resinous, orange-brown.
which it occurs in Mexico. Juniperus californica and Fascicle sheaths initially to 15 mm long, orange-
Quercus turbinella are common; in the cha­parral brown, later 5–8 mm long, dark brown, persistent.
zone many shrubs, e.g. Adenostoma, Ceanothus, Leaves in fascicles of 2–3 (rarely 4 or 5), persisting
Artemisia, Cercocarpus, Rhus, Erio­dictyon, Arcto­ ca. 3 years, straight or curved to twisted, rigid, 8–15
staphylos, and Yucca, dominate. Most of these moun- cm long, 1.1–1.6 mm wide, with serrulate margins,
tains are granitic, but in the south of the ranges more acute-­pungent, bright green or dark green. Stomata
volcanic rock is found. Pinus quadri­folia often grows on all faces of leaves. Pollen cones oblong to cylin-
in cracks among boulders. Annual precipitation is a drical, 1.5–2 cm long, pink to reddish, turning yel-
moderate 300–500 mm, but it is very variable; most lowish brown. Seed cones usually in whorls of 2–5,
of it comes during winter cyclonic storms and there sometimes solitary, on short, stout peduncles, very
is a long dry season from spring through summer. tenacious, serotinous. Mature cones ovoid, nearly
symmetrical or asymmetrical with an oblique base,
(5–)7–12(–15) × (4–)6–10(–12) cm when open. Seed
Conservation
scales parting after a considerable time or only after
IUCN: LC extreme heating, oblong, straight, thick woody. Apo-
physis slightly raised, transversely keeled, or gibbous
especially towards base on one side of asymmetri-
Uses
cal cones, rhombic to weakly pentagonal in outline,
There is no use of this species for timber. It is lo- with radial lines or markings, lustrous light brown.
cally used for firewood. The seeds are edible and are Umbo dorsal, flat or depressed, without a prickle.
harvested to be sold in local markets. Resin may be Seeds obliquely ovoid, (5–)7–10 × 4–6 mm, grey-
tapped on a small scale as well. It is not in cultiva- brown with dark spots. Seed wings 14–20 × 7–10
tion outside a few botanic gardens and other collec- mm, yellowish brown, with grey-brown stripes.
tions; it should be suitable in regions with hot, dry
summers and can endure frost and snow if not for
Distribution
prolonged time and of high altitude provenance.
USA: California, Pacific coast in three localities
from Monterey southward; Mexico: Baja California
Pinus radiata D. Don, Trans. Linn. Soc. London 17: Norte (Cedros Island, Guadalupe Island).
442. 1836. TDWG codes: 76 CAL 79 MXI-GU MXN-BC

Etymology Ecology
The species epithet is said to refer to the radial mark- Pinus radiata is in its natural habitat confined to
ings on the apophyses of some cones. promontories and strips of rocky coast as well as two
offshore islands; it is rarely found more than 11 km
from the sea. The cold ocean water moving down
Vernacular names
the Californian coast substantially influences the lo-
Monterey pine, Radiata pine; pino insigne (Spanish) cal climate, raising humidity, tempering heat from
the sun and causing almost daily summer fogs. On a Pinus radiata D. Don var. radiata. Type: USA:
rare clear day in summer temperatures can quickly California, Monterey, near the coast, J. Coulter
rise to 40 °C, coming down to only 15 °C on the next (type specimen not located, a seed cone).
foggy day. On Guadalupe Island, which lies ca. 250
km offshore surrounded by deep, cold water, fog is
Description
nearly perpetual during the growing season. Rain
fall is erratic, fog drip provides most of the mois- Leaves in fascicles of 3, rarely 2. Seed cones (7–)9–15
ture to the soil and the trees. On the islands the spe- cm long, usually strongly asymmetrical.
cies does not grow below 300 m and ascends to the
summit ridges at 1100 m a.s.l. on Guadalupe Island,
Distribution
but on the mainland it grows from sea level to the 749
base of inland hills and does not exceed 400 m a.s.l. USA: California (Monterey, San Luis Obispo, San
The island populations form pure stands with some Mateo & Santa Cruz Counties).
undergrowth of Quercus tomentella (Guadalupe Is- TDWG codes: 76 CAL
land) or Juniperus californica (Cedros Island). On
the mainland it forms pure stands or mixed stands
Conservation
with Cupressus macrocarpa, Pinus attenuata (both
like P. radiata with serotinous cones adapted to fire), The total area of occupancy (AOO) of Pinus radiata
Arbutus menziesii, and Quercus agrifolia. In the ab- var. radiata has been reduced to about 50% of the
sence of fire, Pseudotsuga menziesii establishes itself, ca. 10,000 ha estimated to have been covered by it
especially in the southernmost population at Año on the arrival of Europeans in California. The cur-
Nuevo, and would become dominant to the exclu- rent main, but still minor threat to this variety is
sion of P. radiata. In stands with Cupressus macro­ forest succession, particularly in the southernmost
carpa there is very little or no undergrowth. population. Pinus radiata var. radiata has extended
it range there in recent decades, but succession to a
forest type that eventually excludes this pine is also
Uses
taking place in the absence of fire. Urbanisation in-
Radiata pine, as it is now commonly called among terferes with the natural fire cycles which favour re-
foresters internationally, is the most widely plant- generation of P. radiata var. radiata. The (genetic)
ed tree species in the world. Its tiny natural relict distinction between natural and semi-natural occur-
stands fall into insignificance to the millions of hec- rence is also blurred by plantings in this and other
tares planted in Australia, New Zealand, Chile, Ar- areas. The accidental introduction of pitch canker in
gentina, Uruguay, South Africa, Kenya, and Spain. the 1990’s (Fusarium circinatum [anamorph] Asco­
Provenance is almost exclusively from mainland mycetes) poses another recent threat.
California. Its spectacularly rapid growth under IUCN: NT
plantation conditions is the main reason for its suc-
cess in commercial forestry and for many of the Pinus radiata D. Don var. binata (Engelm.) Lem-
above mentioned countries it is the most important mon, Handb. W. Amer. Cone-bearers, ed. 3: 42.
timber tree. The wood is rather brittle and coarse 1895. Pinus insignis Douglas ex Loudon var. binata
grained and most suitable for pulp wood, but in Engelm., in Watson, Bot. California 2: 127–128.
many countries where it has been introduced it is 1880; Pinus radiata D. Don subsp. binata (Engelm.)
also put to other uses. These are e.g. construction, E. Murray, Kalmia 13: 24. 1983. Type: Mexico: Baja
carpentry and joinery, veneers, furniture, laminated California Norte, Guadalupe Island, E. J. Palmer s.n.
wood, and crates and boxes. This species has been (holotype MO).
widely planted as a landscape tree in urban areas,
parks and large gardens, where it can grow to huge Pinus muricata D. Don var. cedrosensis J. T. Howell,
size in relatively short time. It is a very suitable tree Leafl. W. Bot. 3: 7. 1941; Pinus radiata D. Don var.
to form a living screen against wind and traffic noise cedrosensis (J. T. Howell) Silba, Phytologia 68: 60.
and tolerates relatively high levels of air pollution. 1990.

2 varieties are recognized:

 Description Description
Leaves in fascicles of 2(–3 on leading shoots). Seed Small trees or shrubs to 3–9 m tall, d.b.h. to 15–40 cm;
cones (5–)6.5–9(–13) cm long, moderately asym- trunk short, contorted, soon branching. Bark thick,
metrical. rough and scaly, exfoliating with thin, scaly plates,
in old trees on lower part of trunk longitudinally
furrowed, grey to blackish grey. Branches spreading
Distribution
or ascending, the ultimate branches stout, assurgent.
Mexico: Baja California Norte (Cedros Island, Shoots rough with prominent, non-decurrent pul-
Guadalupe Island). vini, which leave circular scars after fascicles have
750 TDWG codes: 79 MXI-GU MXN-BC fallen. Cataphylls ca. 5 mm long, subulate, reflexed,
grey. Vegetative buds ovoid to ovoid-cylindric; ter-
minal bud 5–7 mm long; lateral buds shorter, acute,
Conservation
not resinous or sometimes slightly resinous. Fasci-
This variety, confined to two islands, was until very cle sheaths initially ca. 5 mm long, soon caducous,
recently under serious threat from feral goats on not recoiling to form a rosette. Leaves in fascicles of
Guadalupe Island, which is almost uninhabited. 2(–3), persisting 4–5 years, slightly curved to falcate,
It is estimated that only 260 mature trees remain rigid, (2–)3–4.5(–5.5) cm long, 0.8–1.1 mm wide;
there and reproduction is scarce. Total eradication margins entire; apex acute-acuminate; abaxial face
of goats appears to be difficult, but may be achieved dull light green or yellowish green; adaxial face(s)
in the near future as international pressure on the slightly glaucous, with inconspicuous whitish sto-
Mexican government is increasing. Exclosures to matal lines. Stomata on all faces of the leaves. Pol-
keep goats out of some of the most sensitive areas len cones subglobose to ovoid, 4–5 mm long, pink
have been realized with help from outside and this or purplish, turning light yellow. Seed cones solitary
has helped establishment of several hundred pine or sometimes in pairs on 5–8 mm long, slender,
seedlings. ­Pinus radiata var. binata does better on curved peduncles, maturing in two years. Mature
Cedros Island which has (had) no feral goats. cones subglobose or globose when closed, (2–)2.5–4
IUCN: EN (C1) × 3–6 cm when open. Seed scales parting widely;
basal scales remaining connate; all scales loosely
attached to the rachis, irregularly shaped, concavo-
Pinus remota (Little) D. K. Bailey & Hawksw., convex, with incurved margins and 1–2 deep seed
Phytologia 44: 129. 1979. Pinus cembroides Zucc. cavities. Apophysis prominently raised, transversely
var. remota Little, Wrightia 3: 183. 1966. Type: USA: or radially keeled, irregularly rhombic to pentagonal
Texas, Val Verde Co., 48 km N of Del Rio, 21 km in outline; upper margin angular and irregular; col-
S of Loma Alta, on “cedar brake”, E. L. Little & our (lustrous) light brown to reddish brown. Umbo
D. S. Correll 18991 (holotype US). dorsal, flat or indented, with a minute, deciduous
prickle. Seeds obovoid-angular or ellipsoidal, 12–16
Pinus catarinae Rob.-Pass., Bull. Mus. Hist. Nat. × 8–10 mm, light ochraceous with a grey tinge; in-
(Paris), ser. B, Adansonia 1: 70. 1981. tegument very thin, 0.1–0.4 mm; megagametophyte
(‘endosperm’) white. Seed wings vestigial, remnants
remain on the scale when the seeds are dispersed.
Etymology
The species epithet refers to the relatively large dis-
Distribution
tance between leaf fascicles.
Mexico: NE and SE Chihuahua, Coahuila, and ex-
treme W Nuevo León; USA: Texas (Edwards Pla-
Vernacular names
teau, W. Texas along the Rio Grande).
Texas pinyon pine, Nut pine; piñon (Spanish) TDWG codes: 77 TEX 79 MXE-CO MXE-CU MXE-
NL
 Bark on young trees and in crown of older trees thin,
Ecology
orange-brown, on trunk becoming thick and break-
The populations of Pinus remota are almost all ing into irregularly shaped but wide, elongated,
highly disjunct; the species has obviously retreat- scaly, reddish brown plates divided by grey-brown
ed to isolated mountains. Its altitudinal range is fissures. Branches spreading wide, forming a round-
1200–1600(–1850) m a.s.l. (on the Edwards Plateau ed crown. Foliage branches moderately slender, new
in Texas it occurs considerably lower, the type col- shoots to 1 cm thick, light orange-brown or red-
lection is from 450 m). It is restricted to canyons or brown, turning darker brown, rough with pulvini
rocky mountain slopes, often on calcareous soil or from fallen leaf fascicles. Buds ovoid to cylindrical;
limestone rock, on dry sites where Pinyon-Juniper terminal bud to 20 mm long, resinous; cataphylls
woodland is not well developed and the pines often red-brown, fringed, some free at the apex. Leaves 751
remain shrubby. Annual precipitation ranges from in fascicles of 2, held in a basal sheath initially 15–
300–500 mm, but is extremely variable from year 20 mm long, but eroding to basal part of only 5–6
to year. Frost is common in December and January. mm length, persisting 3–4 years, straight or curved,
This species occasionally occurs with P. cembroides 12–18 cm long, slightly twisted, brittle, breaking eas-
and more rarely with P. arizonica var. stormiae, ily, ca. 1.2 mm wide, dark green; margins minutely
common are Juniperus monosperma and J. ashei serrulate; apex abruptly acute; stomata in fine lines
(in the northern part of the range), Quercus, Cerco­ on all surfaces. Pollen cones clustered at base of new
carpus and semi-desert plants e.g. Agave lecheguilla, shoots, spirally arranged, ovoid-oblong, 1.5 cm long,
­Opuntia and Fouquieria splendens. conspicuously red or purple to dark purple, turn-
ing dark brown shortly before they fall. Seed cones
solitary or in pairs, nearly sessile, falling without the
Conservation
short peduncle and often leaving a few basal scales
IUCN: LC behind, 4–6 cm long, narrowly ovoid-conical when
closed, symmetric or slightly asymmetric at base,
opening soon after maturation to a broadly ovoid
Uses
or sub-globose shape with a more or less level base.
Any use of this species (firewood, edible seeds) is Seed scales thin woody, rigid, oblong, dark brown;
probably incidental or at most on a very small scale apophyses slightly raised, transversely keeled, some-
due to its scattered occurence in often remote places. times more pronounced on exposed side of cone
The edible seeds are harvested and sold with those of near base; umbo central, flat or depressed, unarmed
other pinyons. It is not known to be in cultivation or with a minute, deciduous prickle. Seeds ovoid,
outside a few botanic gardens, but should be well 4–5 mm long, slightly flattened, rugose, brown; wing
adapted to gardens with a dry, hot summer. 15–20 mm long, narrow, pale brown.

Distribution
Pinus resinosa Aiton, Hort. Kew. 3: 367. 1789. Type
not designated. E North America, from Newfoundland and West
Virginia westward to Manitoba and Minnesota.
TDWG codes: 71 MAN 72 NBR NFL-NE NSC ONT
Etymology
QUE 74 ILL MIN WIS 75 CNT MAI MAS MIC NWH
The species epithet means ‘resinous’. NWJ NWY PEN VER WVA

Vernacular names Ecology

Red pine, Norway pine; pin rouge (French) Pinus resinosa is a species of pine occupying the
northeastern mixed conifer-deciduous broad-
leaved forest bordering on the boreal conifer forest
Description
at its northern limit. This area was heavily glaciated
Trees to 37 m tall; trunk to 1.5 m d.b.h., straight, during the last Ice Age and the glaciers left huge de-
shorter and with heavy branches in solitary trees. posits of sand, some of which in post glacial time
 was blown into sand dunes. It is to these sands that is the reverse and several cultivars, mainly dwarfed
this pine is largely confined, as here it can success- and shrub forms, have been produced for garden
fully compete with other trees. Another type of use.
habitat for P. resinosa is swamp margins. The species
avoids non-acidic soils and only grows over lime-
stone when there is a leached, acid top layer. It is Pinus ×rhaetica Brügger, Flora 47: 150. 1864. Type:
largely a lowland pine growing at altitudes between Switzerland: Ober-Engadin, K. I. Christensen CH
200 m and 450 m, but ascends to 1300 m a.s.l. in 2–5a (neotype C).
the Appalachian Mountains. In the northern parts
of its range it can form extensive pure stands, but
Etymology
752 more often it is growing together with Pinus banksi­
ana, P. strobus, or both. Populus spp. are common in The epithet refers to the Rhaetica Alps, located in
the sub-boreal zone, further to the south Abies bal­ E Switzerland and W Austria.
samifera, Acer ­rubrum, A. saccharum, Quercus alba,
Q. rubra, ­Fagus grandifolia, and Betula papyrifera
Vernacular names
can come into the forest, especially on more fertile
soils. In mountainous areas, Pinus rigida, P. pun­ None are recorded.
gens, P. virginiana, and Tsuga canadensis are other
conifers growing with P. resinosa. Where this pine is
Description
attaining good size most of the associated tree spe-
cies remain lower, but in some of the broad-leaved This natural hybrid between Pinus mugo Turra and
forest types P. resinosa is only a minor component Pinus sylvestris L. differs from P. mugo in its habit: it
restricted to the poorest sites. This pine is dependent is a tree, rarely a shrub. Its bark on major branches
on fire for its natural reproduction and is therefore a resembles that of P. sylvestris in being reddish brown
seral species on most sites. with papery flakes, but it can also be more like
P. mugo and greyish and persistent. Its seed cones
resemble those of P. mugo, with strongly developed
Conservation
apophyses on one side, and are often more robust
IUCN: LC than those of P. sylvestris.

Uses Taxonomic notes

Red pine is an important timber tree and is widely Hybridization between Pinus mugo and/or its vari-
planted for this purpose in the NE USA and E Can- ous forms and P. sylvestris has been described by
ada. Its other English name, Norway pine, refers to several authors from several localities within the
the homeland of the men who logged this pine in range of P. mugo s. l. and several of these hybrids
Minnesota, where it is the state tree. The pale red- have been named (Christensen, 1987; Vidaković,
dish wood is close-grained and used mostly for 1991). Probably the most common form of this is the
pulpwood and rough construction timbers used in hybrid species listed here and described in detail in
piling, cabin logs, posts, railway sleepers, paving Christensen (1987) and Christensen & Dar (1997).
blocks, crates, and nowadays less frequently mine
timbers. It is also planted for Christmas trees, which
Distribution
use requires pruning the branches to obtain a de-
sired shape and density of foliage. Large tracts of Europe: from Pyrenees to western Carpathians.
dunes have been planted with this pine to stabilize TDWG codes: 11 AUT-AU CZE-CZ GER POL SWI 12
blown sand and the species has been used to form FRA-FR 13 ITA-IT
shelter belts to prevent wind erosion on farmland.
Red pine was introduced to Britain as early as 1756,
Conservation
but is there little used in amenity planting and hor-
ticulture. In the NE USA and E Canada the situation IUCN: LC
Pinus rigida Mill., Gard. Dict., ed. 8: Pinus No. 10.
Distribution
1768. Type not designated.
Extreme SE Canada (Ontario, Quebec); NE and E
USA westwards to Kentucky and Ohio, southwards
Etymology
to South Carolina.
The species epithet means ‘rigid’ or ‘stiff ’ and refers TDWG codes: 72 ONT QUE 75 CNT MAI MAS NWH
to the stiff needles. NWJ NWY OHI PEN RHO VER WVA 78 GEO KTY NCA
SCA VRG TEN
Vernacular names
Ecology
Pitch pine, Hard pine; pin rigide (French) 753
This species occupies the cool maritime and partly
mountainous NE of the USA, where it occurs from
Description
sea level in the north to nearly 1400 m in the south-
Trees to 30 m tall; trunk to 1 m d.b.h., straight or ern Appalachians. It grows on shallow sandy or
curved near base, shorter to 20 m in the northern gravelly soils poor in nutrients, usually well drained,
part of its range. Bark thick and breaking into ir- but sometimes water-logged, as in the swamps of
regularly shaped but elongated, scaly, reddish brown New Jersey. It is a seral species, most commonly
plates divided by deep and long, dark fissures. Ad- associated with oaks (Quercus spp.) or with other
ventitious foliage shoots common on trunk; prima- pines e.g. P. virginiana, P. echinata, or P. pungens. In
ry branches spreading and ascending, often bent to swamp areas it grows with Chamaecyparis thyoides.
one side, poorly self pruning, forming an irregular On some sandy sea shores (e.g. at Cape Cod, Massa-
crown. Foliage branches sparse, assurging, moder- chusetts) a decumbent, wind-swept form 1–5 m tall
ately slender, new shoots 0.5–1 cm thick, yellowish occurs. It is exceptional in its capacity to resprout
green soon turning orange-brown or red-brown, from stumps; small tufts of adventitious foliage ap-
rough with pulvini from fallen leaf fascicles. Buds pear on the trunks even of healthy trees. Its ability to
ovoid to cylindrical; terminal bud 15–20 mm long, regrow foliage after severe damage at any stage in its
resinous; cataphylls appressed, red-brown, fringed; development enables this species to survive fires as
apex cuspidate. Leaves in fascicles of 3, occasionally well as browsing of seedlings and saplings. In areas
2 (adventitious shoots can have dwarf shoots with with regular occurrence of fire many trees are multi-
3–5 leaves), held in a basal sheath 10–15 mm long, stemmed from a basal stool.
persisting 2–3 years, straight or curved, 6–12(–15)
cm long, rigid, twisted, 1.2–1.5(–2) mm wide, green;
Conservation
margins minutely serrulate; apex abruptly acumi-
nate; stomata in fine lines on all surfaces. Pollen IUCN: LC
cones clustered at base of new shoots, spirally ar-
ranged, cylindric, 1.5–2 cm long, yellow. Seed cones
Uses
solitary or in whorls of 2–5, nearly sessile, persistent,
variously serotinous or opening in the second year, The wood of Pitch pine is coarse and resinous, of
3–8 cm long, ovoid-conical when closed, symmetric, small or moderate size and often not straight for any
opening to a broadly ovoid or subglobose shape to considerable length except in planted and well main-
7 cm wide with a level or slightly convex base. Seed tained stands. It is therefore of limited commercial
scales thin woody, rigid, oblong, dull brown with a value, used mainly for rough construction. [‘Pitch
red-brown sealing band; apophyses slightly raised, pine’ in the timber trade may refer to P. palustris in-
more or less rhombic, markedly transversely keeled, stead of this species.] The high resin content makes it
light brown or light red-brown; umbo dorsal and decay resistant, a property which was made use of in
central, low triangular, armed with a slender, curved past ship building and more recently railway sleep-
prickle. Seeds obliquely obovoid, 4–5(–6) mm long, ers. If ‘destructively’ distilled it yielded naval stores
slightly flattened, dark brown, mottled black or near i.e. turpentine and related chemicals, also important
black; wing 15–20 mm long, blackish brown. Perhaps in times of wooden ships. It now finds more use in
the only pine with a (moderate) coppicing capacity. charcoal burning. The adaptibility of this pine to
 poor soil conditions makes it suitable for replanting ovoid-conical when olive green, growing to 10–15
of wasteland. It is of some interest in horti­culture be- (–20) cm long, 7–12 cm wide when closed, opening
cause of its tufts of foliage on the stem and its persist- only slightly after several years to 13 cm max. width.
ent cones, but usually confined to the arboreta and Seed scales oblong, woody and rigid; apophyses
pineta of botanic gardens and private collectors, and strongly developed, thick, conical, with rhombic to
only a few cultivars are known. Its resprouting from irregularly penta­gonal bases, sharply transversely
epicormic buds make it ideal for bonsai ­culture. keeled, (strongly) recurved, smooth, lustrous yellow-
ish brown or grey-brown; umbo triangular, obtuse
and unarmed. Seeds obovoid, 8–12(–15) mm long,
Pinus roxburghii Sarg., Silva N. Amer. 11: 9. 1897. slightly flattened; wing adnate, oblique, 20–25 mm
754 Type: India: Himalaya, [locality not given], W. Rox­ long, 8–10 mm wide, lighter brown than the seed.
burgh B-W 17762 (holotype not located, isotype
B-W).
Distribution
Himalaya, from Pakistan to NE India, Arunachal
Etymology
Pradesh (Assam, Kameng District).
The species name commemorates William Roxburgh TDWG codes: 36 CHT 40 EHM-AP EHM-BH EHM-
(1751–1815), a Scottish botanist working for the East DJ EHM-SI NEP PAK WHM-HP WHM-JK WHM-UT
India Company and Superindentent of the Calcutta
Botanic Garden.
Ecology
Pinus roxburghii is widespread and common in the
Vernacular names
north-south oriented outer valleys of the Himalaya
Chir pine, Long-leaved Indian pine; Chil (Hindi); and its foothills and often forms pure stands espe-
Dhup (Nepalese); Chir, Sula (Pakistan) cially on dry, fire-prone slopes. Mature trees are
relatively fire resistant; regeneration after destruc-
tive fires can be massive and rapid when it acts as
Description
a pioneer species. In prolonged dry seasons it may
Trees to 50–55 m tall, usually not taller than 30 m; drop most of its leaves. It occurs on a variety of
trunk to 3 m d.b.h., usually to 1 m d.b.h., bole substrates, from deep soil to bare rocks. Its altitu-
straight and columnar. Bark becoming thick and dinal range is from 400 m to 2300 m a.s.l., with the
divided by long, deep fissures, breaking into large, highest growing, scattered individuals at 2500 m.
elongated, scaly plates, weathering grey-brown, with Pinus roxburghii is restricted to the monsoon belt
dark brown and purplish hues. Branches self-prun- with summer rains. In its higher altitudinal range
ing, long and spreading to ascending, forming an this pine species is commonly mixed with Cedrus
open, domed crown in mature to old trees. Foliage deodara and Pinus wallichiana, but occurs below
branches slender or stout, pale grey or light brown, the forest zone characterized by species of Abies.
covered with leaf-like brown cataphylls remaining Broad-leaved trees (­angiosperms) are commonly
several years and becoming recurved at their apices. Quercus incana, ­Schima wallichii and Rhododendron
Buds small, ovoid, without resin. Leaves near ends arboreum. Towards its lower limit angiosperms be-
of branchlets, persisting 1–2 years, in fascicles of come more dominant and the pines occur on rocky
3, held together by persistent, 25–30 mm long ba- slopes with a northern or eastern aspect.
sal sheaths, spreading and slightly drooping, 25–30
(–35) cm long, pliant, slender, broadly triangular in
Conservation
cross-section, 1.2–1.7 mm wide; margins minutely
serrulate; leaf colour bright green; apex thinly acute- While forest destruction and logging have reduced
acuminate; stomata in fine lines on all surfaces. Pol- the area of occupancy (AOO) of P. roxburghii, it is
len cones clustered near base of new shoots, spirally still covering extensive areas (an estimated 0.87 mil-
arranged, ovoid-oblong, 13–15 mm long. Seed cones lion ha in India alone) and is therefore not consid-
solitary or in whorls of 2–5 on stout branches, short ered to be threatened with extinction.
pedunculate, persistent, heavy, broadly ovoid or IUCN: LC
 Uses Description
Chir pine is an important pine for resin production Trees to 15–30 m, d.b.h. to 30–60 cm; trunk mono­
in the Himalayan region, especially in NW India. It podial, erect, often curved or contorted. Bark up to
has been the basis of the Indian naval stores indus- 5–6 cm thick on lower part of larger trunks, rough
try, initiated by the Indian Forest Department under and scaly, breaking into large, exfoliating plates di-
British rule in 1888 (Langenheim, 2003) to sup- vided by deep, longitudinal fissures, dark brown,
ply the British Empire with turpentine and related weathering grey. Branches spreading and ascending,
products. Later, it became the main source of these often contorted; crown pyramidal in young trees,
substances for India, but production was falling dra- open and irregular in old trees. Shoots slender, gla-
matically due to poor management of the forests brous and smooth, greyish. Cataphylls ca. 5 mm 755
and destructive tapping methods. In recent years long, fragile, soon deciduous. Vegetative buds ovoid-
these have improved under more rigorous manage- oblong; terminal bud 8–10 × 4–5 mm; lateral buds of
ment. Artificial camphor is the main end product nearly equal size, not resinous. Fascicle sheaths of
derived from resin of P. roxburghii in the region; it young leaves 7–9 mm long, soon recoiling, forming
is also used for medicinal treatments. The wood is a rosette at base of fascicle, finally deciduous before
of importance for railway sleepers after treatment fascicles. Leaves in fascicles of (3–)4–5, persisting
for preservation, and for construction, carpentry 2–3 years on branchlets, straight or slightly reflexed,
and joinery; it is also pulped for the paper industry. lax but not drooping, 6–10 cm long, 0.6–0.8 mm
The bark has a high tannin content (11–14%) and is wide, with irregularly serrulate margins, acute-
used for tanning leather and for staining wood to acuminate, yellowish green or greyish green, weakly
give it an orange colour. The seeds are edible but not glaucous on adaxial faces. Stomata on the two adax-
very good. The needles are used for animal bedding ial faces only. Pollen cones after shoot elongation
and mixed with manure serve as a traditional ferti- forming a 5–7 cm long “spike” leaving shoot base
lizer for agriculture. This species is uncommon in and apex free; cones small, 5 × 3 mm, purplish, turn-
cultivation outside India and Pakistan, but has been ing brown. Seed cones solitary or in whorls of 2–4
introduced to South Africa as a forestry plantation on 15–30 mm long peduncles which fall with cones.
tree. It is sometimes seen as an ornamental tree in Mature cones ovoid to ovoid-conical when closed,
countries around the Mediterranean Sea. Despite ovoid when opened, usually symmetrical, slight-
the fact that in its natural habitat P. roxburghii does ly flattened at base, 10–15 × 6–8.5 cm when open,
not occur in the Himalayan frost zone, some prov- very resinous. Seed scales thin woody, rigid, oblong,
enances could prove to be hardy. concavo-convex, often wider than apo­physis. Apo-
physis prominent, similarly shaped around cone,
transversely keeled, rhombic or pentagonal in out-
Pinus rzedowskii Madrigal & M. Caball., Bol. Técn. line, turning ochraceous or light brown. Umbo dor-
Secr. Agric. Ganad. Subsecr. Forest. Fauna 26: 1. sal, rhombic-pyramidal, transversely keeled, obtuse
1969. Type: Mexico: Michoacan, Coalcomán de or with a minute prickle. Seeds (6–)8(–10) × (4–)5–6
Matamoros, Dos Aguas Forest Station, X. Madrigal mm, brown; wings 20–30(–35) × 8–13 mm, brown,
2202 (holotype INIF). Fig. 247, 248 with darker stripes.

Etymology Distribution
This species was named after the botanist Jerzy Rze- Mexico, W Michoacan (Cerro de Chiqueritas, Cerro
dowski, a student of the Mexican flora. Ocotoso, Puerto del Pinabete).
TDWG codes: 79 MXS-MI
Vernacular names
Ecology
Rzedowski’s pine
The two smaller populations, Cerro de Chique-
ritas and Cerro Ocotoso, are on steep talus of large,
 eroded limestone blocks, near the summits of small Pinus sabiniana Douglas ex D. Don, in Lambert,
mountains in the mainly volcanic Sierra Madre del Descr. Pinus, ed. 8°, 2: p. s.n. inter 144 et 145, t. 80.
Sur. Each have only a few score trees, from old to 1832. Type: USA: California, D. Douglas s.n. 1826
saplings. The larger population is on more level (holotype not located, isotypes GH, K [ex Herb.
ground, in an area known locally as Puerto del Pina- Hort. Soc. London]).
bete, also with limestone boulders, but interspaced
with other substrates. In the two former areas, the
Etymology
trees remain small, less than 15 m tall; in Puerto del
Pinabete trees to 30 m have been found. The alti- This species was named after Sir Edward Sabine,
tudinal range is 2100–2400 m a.s.l. Annual precipi- former President of the Royal Society.
756 tation is ca. 1500 mm, most of it occurs from June
to October. The climate is warm-temperate, with a
Vernacular names
minimum of –5 °C (December) and a maximum of
30 °C (April). Although surrounded by extensive Gray pine, Bull pine, Digger pine (deprecated)
mixed pine forest with species like P. pseudostrobus,
P. herrerae and P. oocarpa, these species do not grow
Description
on the limestone talus. There, Quercus and shrubs,
e.g. Clusia salvinii, form an understorey with Agave Trees to 25 m tall, usually remaining smaller; trunk
and tall herbs. Fires occur frequently, but rejuvena- to 1 m d.b.h., monopodial or forked, straight or
tion seemed good at least at Cerro Chiquerita visited curved. Bark on trunk thick, rough, scaly, divided
by me in 1994. into irregular plates and deep, longitudinal fissures,
brown-grey or blackish grey with reddish brown fis-
sures. Branches of first order long, spreading hori-
Conservation
zontally or ascending, usually persistent along much
No logging seems to take place of this rare pine, of trunk; branches of higher orders sparse, forming
probably because of the inaccessibility of its popula- a broad, irregular and open crown. Shoots slender
tions in remote locations and the poor form of most or stout, rough with prominent, decurrent pulvini
trees from a lumberman’s viewpoint. Mexican bota- from fallen leaf fascicles, pale grey-brown becoming
nists and (regional) foresters are apparently aware of darker brown. Buds ovoid-conical; terminal bud 15–
the uniqueness of this species, but it has not received 25 mm long; lateral buds smaller, all resinous, acute;
a protected status. Studies in population dynamics cataphylls appressed, reddish brown, with paler
related to environmental factors, especially fire and fringed margins. Leaves in fascicles of 3, held by per-
reproduction, are much wanted to ensure proper sistent, initially 20 mm long but later much reduced
management of this evolutionarily interesting pine. (5–7 mm) sheaths, spreading or drooping, persisting
IUCN: EN (D) 3–4 years, (15–)20–28(–32) cm long, pliant, slight-
ly twisted, 1.5 mm wide; margins serrulate; apex
acute-pungent to subulate; leaf colour grey-green;
Uses
stomata in conspicuous lines on all faces of leaves.
No uses are recorded of this species. It is a botani- Pollen cones ovoid to ellipsoid, 10–15 mm long, yel-
cal rarity only present in a few botanic gardens and low becoming orange-brown. Seed cones at base of
private collections. Its altitudinal range implies oc- a subsequent leading shoot, solitary or occasionally
casional light frosts, so it might prove hardy in mild in pairs, on short, persistent peduncles, holding the
regions and as an attractive tree should be worthy of cone to the branch for up to 7 years and retaining a
cultivation, which would also help its conservation few basal scales when falling; mature cones broadly
as a species. ovoid, massive, nearly symmetrical, opening slowly,
17–25 × 15–20 cm when open, more or less flat or
convex at base, extremely resinous. Seed scales thick
woody and rigid, widest towards the apophysis,
with two seed cavities on adaxial side, dull brown;
apophyses very strongly developed, thick woody,
sharply transversely keeled, more or less abruptly
(or gradually) merging into a long, curved umbo, the copious resin to seal drums and baskets. The nu-
up to 20 mm wide, chocolate-brown. Umbos dorsal, tritional value of the seeds can well compete with
elongate, curved, with keeled sides, 10–20 mm long, most other edible pine seeds, yet there is little or no
to12 mm wide at base, ending in a sharp uncinate commercial use at present. The wood, because of
claw. Seeds narrowly obovoid, slightly flattened, its irregular size and shape and high resin content,
15–20 × 7–10 mm, smooth, dark brown; wings short is only used for railroad sleepers, pallets and wood
and wide, ca. 10 mm long, largely ineffective. chips. The needles and twigs yield oils and turpen-
tine. Its irregular, open crown and twisted growth,
usually making several stems, discourages use as an
Distribution
amenity plantation tree; its only interest is in arbo-
USA: California (lower slopes and mountains reta and pineta where the climate is suitable, e.g. in 757
around Central Valley). the south of England, western France, around the
TDWG codes: 76 CAL Mediterranean, or in Australia.

Ecology
Pinus serotina Michx., Fl. Bor. Amer. 2: 205. 1803.
Pinus sabiniana grows in the summer-dry moun- Type not designated.
tains and foothills that encircle the Central Valley
of California, from the edge of the Mojave Desert
Etymology
to the slopes above the Pacific Ocean. Its altitudinal
range is from 50 m to 1800 m a.s.l. Annual precipi- The species epithet (Latin: serotinus = late flowering)
tation varies much within its range, from 250 mm refers to the delayed opening of the cones.
per annum near the desert to 1780 mm at its upper
limits in the Sierra Nevada. It grows inland from the
Vernacular names
coastal fog belt and does not tolerate hard frosts.
Near the coast, it grows in the chapparal zone with Pond pine, Marsh pine, Pocosin pine
ericaceous shrubs which is subject to frequent fires.
On the lower slopes of the Sierra Nevada it grows in
Description
the Upper Sonoran Life Zone, mainly accompanied
by various species of Quercus, as it does at higher Trees to 20 m tall; trunk to 60 cm d.b.h., straight
elevations in the Coast Ranges. Here Pinus coulteri or often crooked. Bark thick and breaking into ir-
can be an associate, in the north of its range it grows regularly shaped, flat but scaly, dark reddish brown
with Juniperus occidentalis. The woodlands with plates divided by deep fissures. Adventitious foliage
P. sabiniana are usually very open, with trees emer- shoots common in crown on largest branches, few
ing from shrubs or standing in areas covered with on trunk; primary branches spreading and ascend-
sparse grasses and herbs. The heavy cones are pre- ing, tortuous, poorly self pruning, forming an irreg-
dated by squirrels and jays, the former can detach ular and thin crown. Foliage branches sparse, stout,
cones from the trees and gnaw through the thick new shoots 1 cm thick, yellowish orange, frequently
scales to obtain the seeds, the latter play a role in glaucous, turning brown, forming numerous buds.
seed dispersal and germination. Buds narrowly ovoid to cylindrical; terminal bud
15–20 mm long, extremely resinous; cataphylls ap-
pressed, red-brown, fringed; apex cuspidate. Leaves
Conservation
in fascicles of 3, occasionally 4 (adventitious shoots
IUCN: LC can have dwarf shoots with 3–5 leaves), held in a
­basal sheath initially 20–25 mm long, reducing to ca.
10 mm, persisting 2–3 years in tufts at the ends of
Uses
branchlets, straight, (13–)15–20(–21) cm long, flex-
Gray pine nowadays has little value as a timber tree; it ible, slightly twisted, 1.3–1.5(–2) mm wide, green;
only played a major part during the California Gold margins minutely serrulate; apex acuminate; stoma-
Rush period due to its proximity to the gold fields. ta in fine lines on all surfaces. Pollen cones clustered
Native American tribes used the seeds for food and at base of new shoots, spirally arranged, cylindric,
 2–3 cm long, yellowish brown. Seed cones solitary a­ ssociated with Taxodium distichum or with other
or in whorls of 2–5, sessile or on a 1 cm long pedun- pines, e.g. P. taeda and P. elliottii. Common broad-
cle, long persistent, variously serotinous or opening leaved trees (angiosperms) belong to the genera
in the second year, (5–)6–10 cm long, ovoid-conical Nyssa, Magnolia, Liriodendron, Persea, and Ilex. Its
when closed, symmetric, opening to a broadly ovoid capability to resprout differs from that of P. rigida in
or subglobose shape to 8 cm wide with a flattened that adventitious shoots appear more in the crown
base. Seed scales thin woody, rigid, oblong, dull than on the trunk, probably because fires tend to be
brown with a dark red-brown sealing band; apo­ crown fires in the wetter forests with water-logged
physes slightly raised, more or less rhombic, trans- forest floors where this species occurs.
versely keeled, light brown or pale red-brown; umbo
758 dorsal and central, small, armed with a short, weak
Conservation
prickle, sometimes unarmed. Seeds obliquely ellip-
soid, 5–6 mm long, slightly flattened, pale brown, or IUCN: LC
mottled dark brown; wing 15–20 mm long.
Uses
Taxonomic notes
The wood of Pond pine is coarse grained and res-
Although this pine forms hybrids with Pinus rigida, inous, while the tree only attains modest size and
as well as with P. taeda, it is generally considered to is often crooked or divided into long branches at
be a distinct species. It is undoubtedly closely relat- two thirds of its height. Its wood is therefore main-
ed to P. rigida, with which it shares the uncommon ly used as pulp wood. Slow growth in the nutrient
character of adventitious shoots. There are sufficient poor acidic swamps makes commercial use unlikely;
independent character differences to separate the however, its growth is considerably better in plan-
two species, although several of these are quantita- tations on well drained sandy soils. It is of limited
tive (to do with measurements) rather than qualita- value in amenity planting and horticulture and at-
tive (absent or present). Distribution and ecology of tempts at forestry plantations in China, South Africa
the two species only overlap marginally. and Zimbabwe have generally had poor results com-
pared to other pine species.
Distribution
SE USA: from New Jersey to Alabama and Florida Pinus sibirica Du Tour, in Déterville, Nouv. Dict.
(coastal plain). Hist. Nat. 18: 18. 1803. Pinus cembra L. var. sibirica
TDWG codes: 75 DEL MRY NWJ 78 ALA FLA GEO (Du Tour) G. Don, in Loudon, Hort. Brit. 1: 387.
NCA SCA VRG 1830; Pinus cembra L. subsp. sibirica (Du Tour)
Krylov, Fl. Altai. Tomsk. 7: 1724. 1914 [& in Fl. Zap.
Sibir. 1: 77. 1927]. Type not designated.
Ecology
Pinus serotina is a lowland pine of wet, swampy ar- Pinus hingganensis H. J. Zhang, Bull. Bot. Res.
eas on the coastal plains of the Atlantic Ocean and North-East. Forest. Inst. 5 (1): 151. 1985; Pinus sibirica
the Gulf of Mexico. It occurs on sandy flats with Du Tour var. hingganensis (H. J. Zhang) Silba, Phy-
a high water table and in swamps. The climate on tologia 68: 61. 1990.
these plains is mild and humid with annual precipi-
tation ranging from 1120 mm to 1420 mm, with the
Etymology
driest months in winter. In so-called pocosins, peat
swamps that have risen above the water table and The species epithet refers to its natural occurrence
drain outwards, P. serotina is common (a local ver- in Siberia.
nacular name is Pocosin pine). Ponds are swamps
in depressions with very poor drainage. In both
Vernacular names
types of swamp the pines form a tall, open canopy,
under which is a dense and varied shrub layer char- Siberian pine, Siberian stone pine; Sibirskii kedr,
acterized by Smilax laurifolia. Pinus serotina is often Kedr (Russian)
 Description Ecology
Trees to 35–40 m tall; trunk to 1.8 m d.b.h. Bark This is a species growing both in lowland areas along
smooth and pale brown on young trees and on the great Siberian rivers like the Ob and the Yenisei
branches, becoming scaly and deeply fissured on Rivers between 100 m and 200 m a.s.l., and in the
trunks of large trees, turning grey. Branches nu- mountains to an altitude of around 2400 m, where it
merous, spreading and assurging, forming a dense, forms dense, pure forests. In the lowlands, it occurs
broadly conical crown. Foliage branches slender to with Pinus sylvestris, Larix gmelinii or L. sibirica and
stout, initially with a dense, pale yellow pubescence, Betula pendula on drier sites that regularly burn and
then glabrous, smooth, yellowish brown or light with Abies sibirica and Picea obovata (‘dark conifer-
brown. Buds ovoid-conical, without resin; cataphylls ous forests’) and Betula sp. on more mesic sites in 759
reddish brown. Leaves in dense tufts towards end the river basins, in bog margins, and uplands. Pinus
of shoots, persiting 2–4 years, in fascicles of 5 held sibirica is one of the major forest-forming conifers
by deciduous basal sheaths of thin, orange-brown in the Siberian taiga and is estimated to cover ca. 45
scales that fall away in the second year, spreading million hectares. It is the slowest growing species in
wide or forward, straight or slightly curved, more or these extensive forests and may live up to 850 years.
less rigid to flexible but not lax, 6–11(–13) cm long, Its edible seeds are harvested by local people as well
triangular in cross-section, not twisted, 1.2–1.7 mm as birds and other animals; the Eurasian Nutcracker
wide; margins minutely serrulate; leaf colour green; (Nucifraga cariocatactes, fam. Corvidae) is almost
stomata in white lines on the two adaxial faces. Pol- entirely responsible for the effective dispersal of the
len cones clustered, spirally arranged, short cylin- seeds by making food caches.
drical, initially reddish turning red-brown. Seed
cones intitially erect, becoming patent, single or in
Conservation
2–3 in whorls on very short peduncles, remaining
closed or opening only slightly at maturity, from IUCN: LC
glaucous green to purplish when growing, resin-
ous, ovoid-conical, usually longer than wide but
Uses
variable, (5–)7–10(–12) cm long, 4–6(–8) cm wide,
dark brown when ripe. Seed scales imbricate, widely The wood of Siberian pine is considered inferior in
cuneate proximally, with 2 deep seed cavities adaxi- quality to that of Scots pine and Siberian spruce, two
ally, soft woody; apophyses broadly rhombic or of the (co)dominant conifers in Russia/Siberia. How-
widely triangular to semi-orbicular, thickened, pi- ever, it still serves numerous uses, such as round-
lose when growing, longitudinally striated when full wood for poles, sawn timber for light construction,
grown and drying to dark brown; umbo terminal, carpentry, furniture, veneers, utensils, boxes, wood
obtuse, slightly upturned, lighter coloured than the carving, and musical instruments. The wood is soft,
rest of the scale. Seeds oblong-obovoid, 10–14 × 5–7 lightweight and rose-coloured with a good texture
mm, slightly ridged across one end but without a for finer applications. Trees are also tapped for resin,
true wing. mainly to produce turpentine. The edible seeds have
a very high fat content (ca. 65%) and contain many
vitamins. In the Altai Mountains harvests may yield
Distribution
200–300 kg of ‘nuts’ or ‘kernels’ (= seeds) per ha.
China: Heilongjiang (Tuqiang), Nei Mongol, Xin- The seeds have to be separated from the cones with a
jiang; Kazakhstan; Mongolia; Siberia; an isolated mechanical cone-thresher as the cone scales do not
population is reported from the Kola Peninsula in open to release the seeds (which in nature is done
NW Russia. by the Eurasian Nutcracker’s strong bill). In Russia
TDWG codes: 14 RUE RUN 30 ALT BRY CTA IRK this species is commonly planted as an amenity tree
KRA TVA WSB YAK 32 KAZ 36 CHI-NM CHM-HJ CHX in large gardens and parks. Few cultivars are known,
37 MON but in Russia several dwarfed growth forms found
 at high altitude or in peat bogs have been described obovoid, slightly flattened, 8–10 × 5 mm, striated
as botanical forms and may also be in cultivation. In black; wing ca. 15 mm long, articulate, with blackish
Europe, the closely related Arolla pine (P. cembra) is stripes.
more commonly planted.
Taxonomic notes
Pinus squamata X. W. Li, Acta Bot. Yunnanica 14 This species was discovered in 1991 and briefly de-
(3): 259. 1992. Type: China: Yunnan, Qiaojia Xian, scribed in Latin and Chinese, with an illustration, in
X. W. Li 91250 (holotype SWFC). 1992. Its affinity is clearly with Pinus bungeana and
P. gerardiana and together these three pines make up
760 the subsection Gerardianae (Section Quinquef­olius)
Etymology
in subgenus Strobus of the pines (Farjon, 2005b:
The species epithet (Latin squamata = ‘furnished 223). It is distinct from both in several characters,
with scales’) refers to the scaly bark of larger trees. e.g. the smooth grey-green bark turning brown on
older trees, the greater number of needles per fas-
cicle and the cones, which are larger than those of
Vernacular names
P. bungeana but smaller than those of P. gerardiana.
qiao jia wu zhen song (Chinese) In the protologue an error occurs regarding the type
citation: J. Q. Pang “zhao-44” (KUN) is called an iso-
type and X. W. Li 91250 (SWFC) is called the holo-
Description
type. Holotype and isotype have to be duplicates of
Trees to 20 m tall; trunk monopodial, erect, to 60 cm the same collection.
d.b.h. (no old trees are known). Bark smooth, hard,
irregularly exfoliating with thin flakes, exposing light
Distribution
yellowish patches, which later turn from grey-green
to variable brown, creating a multi-coloured pattern China: Yunnan (Qiaojia Xian); known from a single
on trunk. First order and highest order branches in locality only.
pseudo-whorls, spreading and assurgent, forming a TDWG codes: 36 CHC-YN
conical to rounded, open crown. Foliage branches
slender, smooth, densely yellow- or grey-brown pu-
Ecology
bescent, or glabrous, pale reddish brown, becoming
grey-brown. Buds ovoid, to 10 mm long, resinous; This extremely rare pine grows at approximately
cataphylls imbricate, triangular-lanceolate, red- 2200 m altitude on a mountain slope with open
brown. Leaves in remote fascicles of 4–5, held by a (disturbed) woodland and grassland with shrubs.
deciduous basal sheath, spreading, rigid or pliant, It is associated with Pinus yunnanensis and various
9–17 cm long, 0.8–1 mm wide, broadly triangular broad-leaved shrubs and trees.
in cross-section; margins serrulate; apex acute; leaf
colour light green with whitish lines; stomata on all
Conservation
surfaces. Leaf anatomy with a uniform epidermis,
3–5 external (marginal) resin ducts and a single This is one of the rarest conifers known, its popu-
vascular bundle. Pollen cones in elongated clusters lation is restricted to a single locality and consists
on new shoots, spirally arranged, axillary to broad of around 36 (20 mature) trees. They occur on a
cataphylls, ovoid-cylindrical, 10–15 mm long. Seed partly deforested mountain on a NW-facing slope,
cones solitary on 1.5–2 cm long peduncles, erect or surrounded by fire-prone grass and scrubland. The
spreading, falling at maturity, ovoid-conical when area has no official conservation status, but the trees
closed, broadly ovoid when open, (7–)8–10 cm long, are now strictly protected. With help from the lo-
5–6 cm wide, turning from green to variable brown. cal foresters, a farmer in the vicinity has successfully
Seed scales oblong-elliptical, woody and rigid, ca. propagated young trees to be eventually planted in
2.7 × 1.8 cm; apophyses thickened, broadly rhombic habitat.
in outline, transversely keeled; umbo dorsal, flat or IUCN: CR (D)
more or less depressed, unarmed. Seeds oblong or
 or nearly entire margins, acute, dark green to glau-
Uses
cous green. Stomata either on all faces or, more
No uses have been recorded; the species is locally commonly, only on adaxial face. Pollen cones ovoid-
propagated for conservation purposes. It may even- oblong to short cylindrical, 6–10 mm long, yellow.
tually appear in the horticultural domain, first in Seed cones solitary or in pairs or sometimes whorls
China and perhaps later abroad. There is substantial of 3–4 on stout, 15–25 mm long peduncles, matur-
interest as this is most probably the rarest species of ing in two seasons. Mature cones variable, mostly
pine in the world. cylindrical or broad cylindrical to ovoid-oblong
when opened, 12–30(–60) × 7–11 cm when open.
Seed scales spreading obliquely to patent; basal
Pinus strobiformis Engelm., in Wislizenus, Mem. scales recurved or reflexed; most scales with curved 761
Tour N. Mexico: 102. 1848. Pinus ayacahuite margins, thick woody, of similar shape around the
Ehrenb. ex Schltdl. subsp. strobiformis (Engelm.) cone but differentiating from base to apex, with one
E. Murray, Kalmia 13: 21. 1983. Type: Mexico: or two deep seed cavities adaxially. Apophysis more
Chihuahua, Cosiquiriachi, F. A. Wislizenus 231 or less cuneate or elongated, thick at the proximal
(holotype MO). end, thinning out and straight, recurved or reflexed
distally, smooth or longitudinally furrowed, often
Pinus strobiformis Engelm. var. carvajalii Silba, Phy- very resinous, light yellowish brown or dark ochra-
tologia 68: 61. 1990. ceous. Umbo terminal, broadly triangular, obtuse.
Pinus ayacahuite Ehrenb. ex Schltdl. var. novogali­ Seeds obovoid, 12–18 × 8–11 mm, reddish brown or
ciana Carvajal, in McVaugh, Fl. Novo-Galiciana 17: brown. Seed wings vestigial to very small, up to half
48. 1992. the length of the seed in some cones.

Etymology Distribution
The species epithet means ‘shaped like strobus’, i.e. it USA: Arizona, New Mexico, rare in Trans-Pecos
is compared with Pinus strobus. Texas; Mexico: in Sonora, Chihuahua, Coahuila,
Nuevo León, Sinaloa, Durango, Jalisco, very local in
Zacatecas and San Luís Potosí.
Vernacular names
TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CO
pino blanco, pinabete, pino nayar (Spanish) MXE-CU MXE-DU MXE-NL MXE-SL mxe-za MXN-
SI MXN-SO MXS-JA
Description
Ecology
Trees to 25–30 m tall, d.b.h. to 1 m; trunk mono­
podial, straight, erect. Bark on trunk thick, rough Pinus strobiformis is a montane pine of mesic sites,
and scaly, fissured, breaking into small, irregu- its altitudinal range is 1900–3500 m a.s.l. It occurs on
lar plates, dark brown, weathering grey. Branches sites with relatively deep, humus-rich though often
spreading horizontally, curved down or assurgent; rocky soils, especially on N-facing slopes or along
crown conical or pyramidal in young trees, be- mountain streams. It grows in small, pure stands
coming more rounded or irregular in older trees. within pine or pine-oak forest, but more commonly
Shoots smooth, puberulent or glabrous, pale red- it is mixed with P. arizonica, P. engelmannii, P. leio­
dish brown, with 5–10 mm long, early deciduous phylla var. chihuahuana, P. durangensis, P. lumholtz­
cataphylls. Vegetative buds ovoid to ovoid-oblong; ii, and/or various species of Quercus and Arbutus. In
terminal bud 8–15 × 5–8 mm; lateral buds smaller, a more mesic forest type, it is associated with Abies
not or only slightly resinous. Fascicle sheaths 20–25 and Pseudotsuga, and at the highest altitudes with
mm long, soon disintegrating and deciduous at end P. hartwegii. The climate in the Sierra Madre Occi-
of the growing season. Leaves in fascicles of 5, very dental is characterized by summer rains (thunder-
rarely 6, persisting 3–5 years, lax, straight or slightly storms) as well as winter precipitation; winter snows
curved proximally or slightly twisted, (5–)7–11(–12) are common at the higher altitudes.
cm long, (0.6–)0.8–1.1(–1.2) mm wide, with ­serrulate
 slightly twisted, deep green or glaucous green, with
Conservation
lines of stomata on the two adaxial faces only; mar-
IUCN: LC gins minutely serrulate; apex acute to acuminate.
Pollen cones ellipsoid to short cylindrical, 10–15
mm long, yellow. Seed cones clustered, usually 2–4
Uses
together, erect at first, becoming pendulous as they
As one of the ‘soft pines’, P. strobiformis, in the ver- grow on 20 mm long peduncles, soon falling after
nacular of N Mexico often equated with P. ayaca­ seed dispersal, (5–)8–20(–25) cm long, sometimes
huite, is sought after by lumbermen as it is considered highly variable in length on the same tree, oblong cy-
to supply wood of good quality for construction and lindrical becoming ellipsoid cylindrical when open,
762 carpentry work. The wood is also considered good then 4–8 cm wide, usually curved but often straight
for making violins. The resin is used medicinally. It and symmetrical. Seed scales thin woody, more or
is rare in horticulture and mostly confined to arbo- less flexible; small scales near cone base imbricate
reta. Difficulties with the correct identification of and straight or incurved; larger scales more or less
the ‘grade’ or ‘complex’ of species, from P. flexilis to straight or with an incurved apohysis, extremely
P. ayacahuite, to which this taxon belongs, means resinous. Apohysis more or less rhombic in outline
that trees that belong here may be grown under an- on central scales, dull pale brown to grey-brown
other name. (lower part of scales purplish brown), narrowing to
an obtuse terminal umbo. Seeds broadly obovoid to
deltoid, flattened, (5–)7–8(–9) mm long, red-brown
Pinus strobus L., Sp. Pl. 2: 1001. 1753. mottled black; wing 20–28 mm long, pale brown.

Etymology Distribution
The species epithet is probably derived from the E North America: from Newfoundland to N Geor-
Greek strobilos = pine cone. gia, westward to Manitoba and Minnesota; S Mexi-
co; Guatemala (highlands).
TDWG codes: 71 MAN 72 NBR NFL-NE NSC ONT
Vernacular names
PEI QUE 74 ILL IOW MIN WIS 75 CNT INI MAI MAS
Eastern white pine, Weymouth pine; pin blanc MIC NWH NWJ NWY OHI PEN RHO VER WVA 78 DEL
(French) GEO KTY MRY NCA SCA TEN VRG 79 MXC-PU MXG-
VC MXS-GR MXS-OA MXT-CI 80 GUA
Description
Ecology
Trees to 67 m tall (most large trees now only to
50 m); trunk to 1.5–2 m d.b.h., growing to a straight Pinus strobus is widely (and disjunctly) distributed
and columnar bole, scarcely tapering in lower half in regions as widely different in climate and topog-
of its length. Bark smooth in young trees, becoming raphy as Newfoundland and Chiapas, Mexico. The
scaly and eventually deeply furrowed, exfoliating in variety strobus is confined to the NE part of the
small or large, rectangular plates, dark grey-brown. species range, where winters are cold and snowy;
Branches whorled, spreading to assurgent, forming var. chiapensis occurs in the wet mountains with
a conical but in old trees rounded crown. Foliage frequent fog in the S part. These populations were
branches slender, smooth, young shoots glabrous once connected, presumably as late as the last Ice
or puberulent with pale brown, downy hairs near Age, when P. strobus and other trees were all pushed
base of pulvini, grey-green or pale reddish brown, southward before the advance of the Laurentide Ice
becoming grey. Buds small; terminal bud to 10 mm Sheet. In the northen part, P. strobus mainly grows in
long, ovoid-cylindrical, acute, light red-brown, the lowland hills around the St. Lawrence River and
slightly resinous. Leaves in fascicles of 5, persisting the Great Lakes, in the Appalachian Mountains to
2–3 years, held in a soon deciduous sheath of light 1200 m a.s.l. In its southern extension it is confined
orange-brown flimsy scales, spreading, 6–10(–12) to much higher altitudes between 800 m and 2000
cm long, straight, slender, flexible, 0.7–1 mm wide, m a.s.l. Annual precipitation varies greatly from area
to area, with lows at around 500 mm and highs in
Distribution
Mexico to 3000 mm. The southern var. chiapensis
experiences no frost, while long and cold winters are E North America: from Newfoundland to N Geor-
the norm in most of the range of var. strobus. Both gia, westward to Minnesota.
varieties are major or minor components of mixed TDWG codes: 72 74 75 78
forests, with other conifers and/or with broad-
leaved trees. There is a similarity of several broad-
Conservation
leaved (angiosperm) tree species in the forests of the
southern Appalachians and the mountains of Ver- The vast resources of timber available to European
acruz and Chiapas, Mexico, but in the colder north colonists from this large pine had been depleted
P. strobus grows with species not common to both towards the end of the 19th century. However, as 763
the northern and southern ranges. regrowth occurred, this has not threatened the con-
tinued existence and occurrence of the species sig-
Conservation nificantly. Hence, while old growth Eastern white
pine is now very rare, under IUCN criteria this vari-
See under varieties.
ety of P. strobus is not under threat.
IUCN: LC
Uses
Eastern white pine was once the most important Pinus strobus L. var. chiapensis Martínez, Anales
timber tree in eastern North America and in colo- Inst. Biol. Univ. Nac. México 11: 81. 1940. Pinus
nial times the British government forbade European chiapensis (Martínez) Andresen, Phytologia 10: 417.
colonists to cut the larger trees (marked with the 1964; Pinus strobus L. subsp. chiapensis (Martínez)
‘broad arrow’) as it wished to reserve these for the E. Murray, Kalmia 12: 23. 1982. Type: Mexico:
British Navy as ship masts. Its fine grained, smooth Chiapas, Ocotepec, M. Martínez s.n. (lectotype
textured wood low in resin makes excellent con- MEXU).
struction timbers, while doors and windows, furni-
ture, and matches are other uses. In the USA and
Description
Canada it is widely planted both for timber and for
urban planting as shelter belts, as well as restoration Seed cones highly variable in length, 6–25 cm.
of areas disturbed by strip mining of coal. Ameri-
cans and Canadians use this pine for Christmas
Taxonomic notes
trees as its foliage can be clipped into shape. As an
ornamental tree it is not very common in Europe, While Mártinez (op. cit.) described and named this
probably due to susceptability to White pine blister taxon as a variety of P. strobus as it has only insignifi-
rust (Cronartium ribicola, Basidiomycota) and poor cant morphological character differences with that
shape in cultivation. Attempts at forestry plantation species, some others have treated it as a species. It
in Britain also failed due to this disease, as well as does not have smaller cones as these are very vari-
to aphid insect predation (Pineus strobus). In the able in size and no discontinuous character states
colder, drier winters of North America it thrives have been found between this variety and var. stro­
better; several cultivars are well known and used in bus. Statistical analysis applied to continuous char-
gardens, especially some of the dwarfed forms. acter states to demonstrate “significant” differences
between the two taxa (Andresen, 1966) is in my view
2 varieties are recognized: an unconvincing approach to the species problem.
Its considerable distance (2400 km) from var. stro­
Pinus strobus L. var. strobus. Type: North America: bus, which can be explained by late Pleistocene cli-
[locality not stated] “5 K Strobus”, P. Kalm LINN matic events, is not a sufficient reason to assign it
1135.10 (lectotype LINN). the rank of species. If the disjunction persists, it may
evolve truly distinct characters and become a full
species eventually.
Description
Seed cones (8–)10–18(–20) cm long.
 brown. Leaves in fascicles of 2, more or less remote,
Distribution
persisting 2–3 years, spreading, held in a short, per-
Mexico: in Guerrero, E Puebla, Veracruz, Oaxaca sistent basal sheath, 4–7(–12) cm long, straight or
and Chiapas; Guatemala: in the departments of El slightly contorted, rigid, 1–2 mm wide, often slightly
Quiche and Huehuetenango. twisted, green or glaucous green; margins minutely
TDWG codes: 79 MXC-PU MXG-VC MXS-GR MXS- serrulate; apex acute-acuminate or mucronate; sto-
OA MXT-CI 80 GUA mata in fine grey-white lines on all surfaces. Pollen
cones short cylindrical, elongating to 2–2.5 cm, yel-
low. Seed cones solitary or in whorls of 2–5, short
Conservation
pedunculate, usually falling soon after seed disper-
764 Although a valuable timber tree logged locally, the sal, ovoid-conical when closed, near-symmetrical,
main reason for its decline is deforestation and/or (2–)3–7 cm long and 2–3 cm wide, opening to 4–5
forest degradation. cm wide. Seed scales thin woody, rigid, oblong,
IUCN: VU (B1+2b) spreading wide except basal scales, chestnut brown;
apophyses nearly flat to slightly raised, transversely
keeled, more or less rugose, ripening from green
Pinus sylvestris L., Sp. Pl. 2: 1001. 1753. to pale grey-brown or reddish brown; umbo small,
obtuse, unarmed or sometimes with a small, brittle
prickle. Seeds ovoid-oblong, 3–5 mm long, grey-
Etymology
brown with dark specks, or dark brown entirely;
The Latin species epithet sylvestris means ‘growing wing 10–15 mm long, 5–6 mm wide, light brown or
in the forest’. grey-brown.

Vernacular names Taxonomic notes

Scots pine; pin sylvestre (French); Gemeine Kiefer As can be expected, this is a variable species and
(German); furu (Norwegian); tall (Swedish); Sosna more than 50 botanical varieties or subspecies have
zwyczajna (Polish); Sosna lesnaya, Sosna obikno- been named and described. Only three (including
vennaya (Russian) var. sylvestris) are recognized here.

Description Distribution
Trees to 35(–40) m tall (most trees in natural habitats Eurasia: from N Spain and Scotland in the west to
not taller than 25 m); trunk to 1.5 m d.b.h., straight the Russian Far East, from Lapland in the north to
or slightly crooked, sometimes forked in crown. Turkey in the south.
Bark thin and papery, flaking, orange-brown on TDWG codes: 10 FIN GRB NOR SWE 11 AUT-AU
young trees and in the crown of older trees, becom- AUT-LI BGM-BE BGM-LU CZE-CZ CZE-SK GER HUN
ing thick and scaly on lower trunk, breaking into POL SWI 12 FRA-FR SPA-AN SPA-SP 13 ALB BUL
irregular plates, purplish brown weathering grey, GRC ITA-IT ITA-SM ROM YUG-BH YUG-CR YUG-
flaking in small or large chips. Branches spreading KO YUG-MA YUG-MN YUG-SE YUG-SL 14 BLR BLT-
or curving down to pendulous, contorted, form- ES BLT-KA BLT-LA BLT-LI KRY RUC RUE RUN RUS
ing domed or irregular, open crowns in most cases. RUW UKR-MO UKR-UK 30 ALT BRY CTA IRK KRA
Foliage branches slender, spreading or pendulous TVA WSB YAK 31 AMU KHA 32 KAZ 33 TCS 34 TUR
in lower crown, becoming rough with pulvini af- 36 CHI-NM CHM-HJ CHM-JL 37 MON
ter shedding leaves, sometimes remaining smooth,
glabrous, new shoots yellowish green, grey-green or
Ecology
pale yellow, turning light brown to grey-brown or
grey. Buds ovoid or ellipsoid, acute; terminal bud to Accross its enormous range Pinus sylvestris grows
15 mm long; lateral buds smaller, all without resin naturally in a variety of habitats, the common de-
or slightly resinous; cataphylls red-brown or pale nominator of which is deficiency of nutrients in the
soil. Thus on the Atlantic seaboard with high lev- mixed woodland now managed as ‘multiple use’ or
els of precipitation it occupies ancient igneous or even nature reserves, the invasivenes onto Calluna
­metamorphic rocks with little or no soil in Scotland heathland is seen as a menace to biodiversity and
and Norway up to 70° N, while south of the Baltic an ancient semi-natural landscape. In horticulture a
Sea it grows on podzolized glacial sands left after the large number of cultivars is known, including dwarf
Ice Age. In the central Alps it is restricted to the dri- forms from witches brooms; the species is being
er slopes and valleys below other conifers like Larix planted as an amenity tree in many countries.
and Picea, while in the Caucasus it ascends to 2600
m on rocky outcrops and scree. In much of Siberia 3 varieties are recognized:
it occupies the drier sites, but in Scandinavia and
NE ­Europe it often borders acidic peat bogs. In the Pinus sylvestris L. var. sylvestris. Type: [“Habitat 765
steppes of Russia and Mongolia it occurs only along in Europae borealis sylvis glareosis”], Illustration
stream courses. Pinus sylvestris most commonly “Pinus sylvestris” in Daléchamps, Hist. General. Pl.
forms open pine forests and woodlands but in many 1: 45. 1587. Fig. 249
areas it is associated with conifers like Picea, Larix,
Juniperus and with broad-leaved trees, especially Pinus sylvestris L. var. scotica Beissn., in Jaeger &
Betula spp. and Populus tremula. In old growth Beissner, Ziergehölze, ed. 2: 488. 1884.
stands there is often a well developed ground cover
of Vaccinium spp. or Empetrum nigrum in Atlantic
Description
regions, and such pine forests are rich in mycor-
rhizal fungi. Young shoots usually rough with pulvini, light
brown; bud scales (cataphylls) red-brown. Leaves
5–7(–8) cm long. Seed cones variable, but up to 7 cm
Uses
long.
Scots pine is an important timber tree, but most of
the production goes to the paper industry. In the
Taxonomic notes
past it was more often put to use as mining props
and for interior construction; such uses are still There are no morphological or anatomical charac-
common in E Europe. Most of the ‘pine’ used for ters known to clearly distinguish the disjunct Scot-
furniture in W Europe is actually spruce (Picea tish population from those of northern Europe, and
abies), which has a smoother grain and is less res- consequently the ‘Caledonian pine’ is here treated as
inous, but often has more and darker ‘knots’, which Pinus sylvestris var. sylvestris.
are the discarded lower branches on the trunks of
densely planted trees. Other uses of Scots pine wood
Distribution
are (or were) street paving blocks, railway sleepers,
fencing, crates, pallets, boxes, laminated wood, par- Eurasia: from N Spain and Scotland in the west to
ticleboard, fibreboard, and various wood-based ma- the Russian Far East, from Lapland in the north to
terials. In Russia and Scandinavia resin is extracted Turkey in the south.
by ‘destructive distillation’ from the stumps and TDWG codes: 10 FIN GRB NOR SWE 11 AUT-AU
roots of felled trees to produce ‘Stockholm tar’. In AUT-LI CZE-CZ CZE-SK GER HUN POL SWI 12 FRA-
much of western Europe it is a widely planted forest- FR SPA-AN SPA-SP 13 ITA-IT ITA-SM ROM YUG-BH
ry tree for timber; it was introduced in the USA for YUG-CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-
similar purposes and for growing as Christmas trees. SL 14 BLR BLT-ES BLT-KA BLT-LA BLT-LI RUC RUE
Scots pine is or was also used to stabilize dunes, but RUN RUW UKR-MO UKR-UK 30 ALT BRY CTA IRK
not those close to the sea as it is not very resistant to KRA TVA WSB YAK 31 AMU KHA 32 KAZ 36 CHM-HJ
salt-laden winds. In Belgium, the Netherlands, Ger- CHM-JL 37 MON
many, and Denmark such plantations have led to
massive spontaneous spread of pines onto heathland
Conservation
and the last remaining inland dune areas, and while
the old plantations have in many places matured to IUCN: LC
 Pinus sylvestris L. var. hamata Steven, Bull. Soc.
Conservation
Imp. Naturalistes Moscou 11: 52. 1838. Pinus syl­
vestris L. subsp. hamata (Steven) Fomin, Vestn. IUCN: LC
Tiflissk. Bot. Sada (Monit. Jard. Bot. Tiflis) 34: 20.
1914; Pinus hamata (Steven) Sosn., in Grossheim
et al., Fl. Tiflisa 1: 11. 1925, non Roezl. (1857); Pinus Pinus tabuliformis Carrière, Traité Gén. Conif., ed.
sosnowskyi Nakai, Chôsen Sanrin Kaihô 167: 32–33. 2, 1: 510. 1867 [“tabulaeformis”].
1939. Type: Caucasus Mts.: H. Wittmann s.n. (lecto-
type H No. 1002536, right hand specimen).
Etymology
766 Pinus kochiana Klotzsch ex K. Koch, Linnaea 22: 296. The species epithet may refer to a common use of
1849; Pinus sylvestris L. subsp. kochiana (Klotzsch ex the wood and is derived from the Latin tabula =
K. Koch) Eliçin, Istanbul Univ. Orman Fak. Dergisi, plank, and formare = to form or make.
ser. A, 20 (2): 289. 1971.
Vernacular names
Description Chinese red pine; you song (Chinese)
Young shoots pale yellow, turning grey. Leaves 2–7
Description
cm long, glaucescent; apex mucronate. Seed cones
small, 2–5.5 cm long. Trees to 25(–30) m tall; trunk to 1.2 m d.b.h., mo-
nopodial, sometimes forked in or below crown.
Bark scaly, fissured, breaking into large irregular
Distribution
plates, greyish brown weathering grey, flaking in
Ukraine: Krym [Crimea]; Caucasus; Turkey. small or large chips. Branches spreading or curving
TDWG codes: 14 KRY 33 NCS TCS 34 TUR down, contorted, forming a domed or flat-topped
crown. Foliage branches stout, becoming rough with
pulvini after shedding leaves, glabrous, new shoots
Conservation
yellowish brown, pale brown or slightly glaucous,
IUCN: LC turning light brown to grey-brown or grey. Buds
oblong, acute; terminal bud to 20 mm long; lateral
Pinus sylvestris L. var. mongolica Litv., Sched. buds smaller, all slightly resinous; cataphylls ap-
Herb. Fl. Ross. 5: 160. 1905 [“mongholica”]. Type: pressed, pale brown. Leaves in fascicles of 2, some-
NE China: [“Mongholia, … in arenosis pr. stat. viae times 3, more or less remote, persisting 2–3 years,
ferreae Charchonte”], D. Litvinov 1599 (lectotype spreading, held in a 10–15 mm long, persistent ba-
LE). sal sheath, 6–15 cm long, straight or curved, rigid,
1–1.5 mm wide, often slightly twisted, dark green;
Pinus sylvestris L. var. manguiensis S. Y. Li & Adair, margins minutely serrulate; apex acute or acumi-
Sida 16 (1): 184. 1994. nate; stomata in fine lines on all surfaces. Pollen
cones short cylindrical, ca. 2 cm long, yellow. Seed
cones solitary or in pairs, short pedunculate, usu-
Description
ally persisting a few years but opening soon, ovoid
Young shoots smooth, grey-green; bud scales (cata- when closed, near-symmetrical, 5–8.5 cm long and
phylls) pale brown or pale yellowish brown. Leaves 3.5–5 cm wide, opening to 5–7.5 cm wide. Seed scales
4–12 cm long. woody, rigid, oblong to obovate, spreading wide ex-
cept basal scales, often recurving, brown; apophyses
prominently raised, variously shaped, transversely
Distribution
keeled, ripening from green to yellowish brown or
Russian Federation: E Siberia, around Lake Baikal; pale brown; umbo pyramidal, armed with a sharp,
N Mongolia (very rare); China: Nei Monggol, N curved prickle. Seeds ovoid-oblong, 6–8 × 4–5 mm,
Hinggan Range. pale brown, mottled; wing 12–18 mm long, 5–7 mm
TDWG codes: 30 BRY CTA IRK 36 CHI-NM 37 MON wide, light brown or grey-brown.
 Distribution Description
China: from Jilin and Liaoning in the NE to Yunnan Bark grey or brown-grey, breaking into large plates.
in the SW and from Shandong in the E to Qinghai New shoots yellowish or pale brown, not glaucous.
and Sichuan in the W. Leaves rigid, 1.2–1.5 mm wide.
TDWG codes: 36 CHC-CQ CHC-SC CHC-YN CHI-
NM CHI-NX CHM-LN CHN-BJ CHN-GS CHN-HB
Distribution
CHN-SA CHN-SD CHN-SX CHN-TJ CHQ CHS-HN
China: from Jilin and Liaoning in the NE to Yunnan
in the SW and from Shandong in the E to Qinghai
Ecology
and Sichuan in the W. 767
Pinus tabuliformis is most common at middle eleva- TDWG codes: 36 CHC-SC CHI-NM CHI-NX CHN-BJ
tions in the hills and mountains of NE and Central CHN-GS CHN-HB CHN-SA CHN-SD CHN-SX CHN-TJ
China, but occurs from 100 m to 2600 m a.s.l. It pre- CHS-HN
fers dry, sunny slopes and hills where competition
from broad-leaved trees is less severe as the woods
Conservation
are more open and lower than in more mesic sites. It
is also a pioneer in secondary vegetation and is there IUCN: LC
commonly mixed with deciduous shrubs and trees,
in later stages of the succession often giving way to Pinus tabuliformis Carrière var. mukdensis (Uyeki
these angiosperms. Management of forests often has ex Nakai) Uyeki, J. Chôsen Nat. Hist. Soc. 3: 45.
arrested this succession in favour of the pines, which 1925. Pinus mukdensis Uyeki ex Nakai, Bot. Mag.
are of economic importance as a forestry tree. (Tokyo) 33: 195. 1919; Pinus tabuliformis Carrière
subsp. mukdensis (Uyeki ex Nakai) Businský, Acta
Pruhoniciana 68: 26. 1999. Type: North Korea:
Uses
[locality not stated], H. Uyeki 2350 (holotype TI).
Chinese red pine is an important forestry tree yield-
ing timber for construction; its wood is hard and
Description
strong with a straight grain. The wood is used for
mining props, railway sleepers, to build wooden Bark dark grey, longitudinally or irregularly fis-
bridges and carts or wagons, to make tools, and for sured. Leaves 1–1.2 mm wide. Seed cones 4–8.5 cm
artificial fibres. The resin is extracted from the bark long and to 7 cm wide when opened.
and leaves and produces turpentine; another prod-
uct of these parts is tannin used to make leather
Distribution
from hides. Essential oils distilled from the leaves
and the pollen are used in traditional Chinese medi- NE China: Jilin (?), Liaoning; North Korea.
cine. In China, this species is commonly planted in TDWG codes: 36 CHM-LN 38 KOR-NK
plantations for forestry purposes and occasionally
as an ornamental tree. Elsewhere it is uncommon,
Conservation
even in Japan, despite its suitability as an amenity or
forestry tree. While the existence of a few botanical IUCN: LC
varieties, some or all of which may also be in culti-
vation, indicates its potential, no cultivars have been Pinus tabuliformis Carrière var. umbraculifera
selected and named. T. N. Liou & Q. L. Wang, in Liou, Ill. Fl. Lign. Pl. NE
China: 97, 548. 1958. Type: China: Hebei, Anshan
3 varieties are recognized:
Shi, Chin. coll. n.v. (holotype PE?).
Pinus tabuliformis Carrière var. tabuliformis. Type
not designated. Description
Pinus tabuliformis Carrière var. brevifolia S. Y. Wang Trunk monopodial only towards base, usually
& C. L. Chang, Fl. Henan: 1: 135. 1981. forked from below the crown. New shoots yellowish
 or pale brown, not or only slightly glaucous. Leaves (sometimes more) on short peduncles, opening and
rigid, 1.2–1.5 mm wide. falling soon, variable in size, (5–)6–12(–15) cm long,
narrowly conical to oblong when closed, mostly
symmetrical, opening to 6–9 cm wide, becoming
Distribution
more ovoid with a tapering, sometimes oblique
China: Hebei (Central Liaoning, Anshan Shi). base. Seed scales thin woody, oblong, nearly flat,
TDWG codes: 36 CHN-HB dull brown; apophyses slightly raised, those on basal
scales more gibbous, more or less rhombic or with
a rounded upper margin, transversely keeled, often
Conservation
radially striated or wrinkled, dull pale brown; umbo
768 IUCN: LC dorsal and central, pyramidal or recurved, terminat-
ing in a strong, sharp prickle. Seeds obovoid, slightly
flattened, 5–6 mm long, red-brown, often with nu-
Pinus taeda L., Sp. Pl. 2: 1000. 1753. Type: USA: merous black specks; wing 16–20 mm long.
[“Habitat in Virginiae, Canadae paludosis”],
J. Clayton 496 (lectotype BM).
Distribution
SE USA: from Delaware and New Jersey to central
Etymology
Florida and E Texas.
The species epithet is a Latin word for ‘pine tree’ or TDWG codes: 74 OKL 75 DEL MRY 77 TEX 78 ALA
its wood; its other meaning was a torch. ARK DEL FLA GEO KTY LOU MRY MSI NCA SCA TEN
VRG WDC
Vernacular names
Ecology
Loblolly pine, Southern pine
Pinus taeda is widely distributed on the Atlantic
Coastal Plain and extends into the plateaus and
Description
foothills around the southern Appalachians to ca.
Trees to 45 m tall (usually to ca. 30 m); trunk to 1.6 700 m a.s.l. but avoids the Mississippi floodplain.
m d.b.h., usually a straight bole. Bark thin, slightly The climate is warm-temperate and moist, with
scaly, orange-red in crown and upper bole, becom- mild winters and long, hot summers; annual pre-
ing thicker, breaking into more or less square, scaly cipitation is between 1000 and 1500 mm. This pine
plates on large trunks, darker red-brown. Branches forms extensive stands on low, sandy knolls in the
spreading wide, self-pruning leaving a clear bole, ‘prairie swamps’ along the Gulf of Mexico; in the
forming a broadly conical or domed crown. Foliage inland parts of the coastal plain it occurs on river
branches moderately slender, 0.5–1 cm thick, rough floodplains and old river terraces with deep, rela-
with pulvini from fallen leaf fascicles, glabrous, tively dry sandy or loamy soils. This species can
prominently ridged, orange-brown to reddish form pure stands resulting from pioneer invasions
brown, becoming darker with age. Buds cylindrical, after forest disturbance or onto abandoned fields, or
acute, 10–15(–20) mm long, slightly resinous; cata- in mixed pine-dominated forests with several oth-
phylls pale red-brown; margins fringed white; apex er species of Pinus. It is also a component of forest
reflexed. Leaves in remote fascicles of (2–)3, held by types dominated by broad-leaved trees, especially
persistent, 15–25 mm long basal sheaths, remain- species of oak (Quercus spp.) as well as Acer rubrum,
ing 3 years on branchlet, spreading, straight, (10–) Liriodendron tulipifera, Fagus grandifolia, Fraxinus
12–20(–23) cm long, rigid but more or less pliant, spp., and Diospyros virginiana in upland sites. In
slightly twisted, 1.5–2 mm wide, lustrous light green; the coastal swamplands Magnolia grandiflora, Nyssa
margins minutely serrulate; apex acute to abruptly aquatica, Quercus michauxii, Carya aquatica, and
subulate; stomata in lines on all surfaces. Pollen Ulmus americana are common associates of P. ­taeda,
cones cylindrical, 2–4 cm long, 7–10 mm wide, radi- together with other pines and an undergrowth of
ally spreading, yellow. Seed cones solitary or in pairs shrubs and palmettos (dwarf palms).
 stumps on trunk; higher order branches assurgent
Conservation
and densely crowded, forming flat-topped or domed
IUCN: LC crowns in natural habitat. Foliage branches glabrous,
more or less smooth, light brown. Buds ovoid-coni-
cal, 10–15 mm long, 5–7 mm wide, slightly resinous;
Uses
cataphylls appressed, orange or rusty brown. Leaves
Loblolly pine is commercially the most important in fascicles of 2, held by a persistent, slender basal
pine of the S United States, where it makes up over sheath 10–15 mm long, straight or slightly curved,
half of the standing pine volume. It is much used (5–)10–20(–22) cm long, slender, pliant, slightly
in plantation forestry and is a fast grower. Its sawn twisted, 0.7–1 mm wide; margins serrulate; apex
wood properties are not of sufficient quality to be acute; stomata in fine lines on all surfaces. Pollen 769
used in high grade construction and manufacture, cones clustered, spirally arranged, short cylindrical,
and its fast growth and great volume is consequent- 1.5–2 cm long, yellow, becoming yellowish brown.
ly put mainly to the wood pulp industry for paper Seed cones solitary or sometimes in pairs, persistent
and other long-fibre products. In urban settings the on short peduncles, spreading or slightly reflexed,
species finds a use as shelterbelt trees and for soil (3–)4–9(–10) cm long, more or less asymmetrical,
stabilization, again thanks to its rapid growth. This narrowly ovoid when closed, widening to 2.5–5 cm
capacity for quick volume increase has also been when open. Seed scales thin woody, rigid, oblong,
reason to investigate its suitability as a biomass pro- ca, 2.5 × 1.3 cm at mid cone (in larger cones), straight
ducer for the generation of energy. Plantations for when spreading, dull brown; apophyses rhombic in
this purpose are now being exploited and its use outline or with rounded upper margin, nearly flat to
may well increase in future. Loblolly pine has also slightly raised, transversely keeled, slightly rugose,
been introduced in many countries and is grown in lustrous brown; umbo broadly ellipsoid, flat, armed
forestry plantations on a large scale in South Africa, with a minute, often deciduous prickle or unarmed.
Brazil, China, Australia, and New Zealand. This spe- Seeds ellipsoid-ovoid, 5–6 mm long, slightly flat-
cies is little used in horticulture (except the use of tened; wing 15–20 mm long, persistent.
leaf litter as a mulch); more northern provenances
may well be hardy to light frosts. It has a preponder-
Taxonomic notes
ance to invasiveness.
Pinus taiwanensis is similar to P. hwangshanensis,
P. luchuensis and P. densiflora and these species are
Pinus taiwanensis Hayata, J. Coll. Sci. Imp. Univ. closely related according to phylogenetic analy-
Tokyo 30 (1): 307. 1911. ses using DNA sequence data. Recently, Businský
(2003) revisited the morphology of some of these
pines and separated some trees in Taiwan as a new
Etymology
species, ­Pinus fragilissima. Most of the characters
The species epithet denotes its origin, the island of evaluated are either similar to those of P. taiwanen­
Taiwan. sis, or they show overlapping states. The seed scales
are described as “thin” in the formal description and
elsewhere as “fragile” but these are difficult to quan-
Vernacular names
tify and may be attributes of the other taxa as well.
Taiwan black pine, Formosa pine; tai wan song What remains of this are somewhat longer leaves and
(Chinese) only slightly longer seed cones. Species are expected
to show distinct character state differences, without
overlapping states. This taxon should be given the
Description
status of a variety only, as will be done below.
Trees to 45(–50) m tall; trunk to 1.2 m d.b.h, bole
straight. Bark on trunk rough and scaly, breaking into
Distribution
large plates and deep fissures, grey-brown to dark
grey. First order branches long and spreading, slow- Taiwan.
ly and sometimes only partly self-pruning, leaving TDWG codes: 38 TAI
 Pinus taiwanensis Hayata var. fragilissima
Ecology
(Businský) Farjon, comb. et stat. nov. Basionym:
Pinus taiwanensis grows in the mountains or along Pinus fragilissima Businský, Novon 13 (3): 282. 2003.
the mountainous coast; in the mountains of the in- Type: Taiwan: Taitung Co., 1 km N of Wulu along
terior of Taiwan it occurs from 800 m to 3000 m Southern Cross-Island Hwy., R. Businský 32172
a.s.l., with exceptions to 3400 m where it is severely (holotype PR).
stunted. On the coastal slopes it comes down to 600
m a.s.l. Due to this altitudinal range, it occurs in
Description
different forest zones from warm temperate to sub-
alpine, but at the lower and middle elevations it is Leaves (12–)16–20(–22) cm long. Seed cones (5–)6–
770 mostly restricted to open spaces, exposed ridges and 9(–10) cm long; seed scales often somewhat thinner
places with sandy, acidic and nutrient-poor soils. It than in var. taiwanensis.
is associated with various species of oak (Castanop­
sis, Quercus), forming pine-oak woodland in such
Distribution
places.
Taiwan (Taitung Co., Kuan Shan massif).
TDWG codes: 38 TAI
Uses
Taiwan black pine has good quality timber with
Conservation
suitable strength for construction, e.g. in buildings
and wooden bridges, and use as railway sleepers and IUCN: DD
mine props. Its wood is also used for building fences
and gates, crates and boxes, panelling, flooring, fur-
niture making, industrial and domestic woodware, Pinus tecunumanii Eguiluz & J. P. Perry, Revista Ci.
tools, plywood, fibreboard and wood pulp. In horti- Forest. 8: 4. 1983. Pinus patula Schiede ex Schltdl.
culture it is rare except for the use for bonsai grow- & Cham. subsp. tecunumanii (Eguiluz & J. P. Perry)
ing, which is very popular in E Asia. Styles, F. A. O. Forest Genet. Resources Inform. 13:
50. 1984. Type: Guatemala: Baja Verapaz, Sierra
2 varieties are recognized: de Chuacus, San Jéronimo, T. Eguiluz 2 (holotype
GH).
Pinus taiwanensis Hayata var. taiwanensis. Type:
Taiwan: Central Mts., T. Kawakami & U. Mori 2097 Pinus oocarpa Schiede ex Schltdl. var. ochoterenae
(lectotype TI). Fig. 250 Martínez, Anales Inst. Biol. Univ. Nac. México 11:
65. 1940.
Description
Etymology
Leaves (5–)10–15(–17) cm long. Seed cones (3–)4–
6(–8) cm long. This species was named after Tecun Uman, a leader
of the Quiche Indians who was killed during the
Spanish conquest of the Central American Isthmus.
Distribution
Taiwan.
Vernacular names
TDWG codes: 38 TAI
Schwerdtfeger’s pine; pino tecun uman (Spanish)
Conservation
Description
IUCN: LC
Trees to 50–55 m tall, d.b.h. to 1.2–1.4 m; trunk mo-
nopodial, erect, straight. Bark on lower part of trunk
ca. 5 cm thick, grey-brown, above 3–4 m thinning
out, reddish brown. Branches spreading or slightly
ascending, the higher order branches flexible but pine stands with grasses, ­Pteridium aqui­linum, Ru­
not pendulous. Shoots rough, reddish brown, often bus, Calliandra, and Leucaena are predominant as
glaucous. Cataphylls subulate, erose-ciliate at mar- long as the disturbances do not lead to further deg-
gins, scarious, brown. Vegetative buds oval-oblong radation. In less disturbed areas, mostly at higher al-
to cylindrical; terminal bud 15–20 mm long; lateral titudes, P. tecunumanii is often associated with other
buds smaller, not resinous. Fascicle sheaths initially pines, such as P. oocarpa, P. maximinoi, and P. pseudo­
up to 25 mm long, orange-brown, in mature fas- strobus, and at the more mesic sites P. ­ayacahuite and
cicles reduced to 12–18 mm, grey-brown. Leaves P. strobus var. chiapensis. Abies ­guatemalensis and
in fascicles of 4(3–5), persisting 2–3 years, lax and Cupressus lusitanica are other conifers on these high
drooping but not pendant, (14–)16–18(–25) cm long, mountain ridges. On the Atlantic slopes in Chiapas a
0.7–1(–1.3) mm wide, serrulate at margins, acute, mixed angiosperm forest with Liquid­ambar, Magno­ 771
bright green. Stomata on all faces of the leaves. Pol- lia, Clethra, Carpinus, Symplocos, Quercus and many
len cones ovoid-oblong to cylindrical, 1.5–2 cm × other species predominates, and Pinus tecunumanii
5–6 mm when shedding pollen, yellowish. Seed and other pines occur either on poorer sites or at an
cones subterminal, in whorls of 2–4, rarely solitary, earlier stage in a sere leading back to dominance of
on (15–)20–25 mm long peduncles, semi-serotinous, broad-leaved trees.
persisting 1–3 years after shedding seeds, falling with
peduncles. Mature cones ovoid to broadly ovoid,
Conservation
nearly symmetrical or asymmetrical, (3.5–)4–7(–7.5)
× (3–)3.5–6 cm when open. Seed scales thin woody, While occasionally still abundant and of very tall
parting usually within 1–2 years after maturity, ob- stature, this pine is now usually scattered in small,
long, straight or slightly curved. Apophysis raised, disjunct populations and has been depleted by over-
transversely keeled, on proximal scales more or less exploitation and forest clearing, especially at lower
gibbous, striate, dull light brown or slightly lustrous. elevations, to such an extend that many of these
Umbo dorsal, flat or slightly raised, with a minute, populations are now vulnerable to extinction (Dvo-
deciduous prickle, grey. Seeds obliquely ovoid, 4–7 rak & Donahue, 1992).
× 2–4 mm, dark grey-brown, with blackish dots, or IUCN: VU (A2c)
blackish grey; wings articulate, 10–13 × 4–8 mm,
grey-brown.
Uses
Pinus tecunumanii is an important timber tree
Distribution
in Central America, where it can grow a straight
S. Mexico: Oaxaca, Chiapas; Belize; Guatemala; bole with large dimensions. It is largely exploited
Honduras; El Salvador; Nicaragua. for sawn timber and other local wood products in
TDWG codes: 79 MXS-OA MXT-CI 80 BLZ ELS GUA its native range; potentials for wood pulp produc-
HON NIC tion are considered to be high if it was to be grown
extensively in plantations. This taxon has received
considerable interest from foresters as a species for
Ecology
potential plantation forestry to be introduced in
The altitudinal range of this species is considerable: tropical countries. A comprehensive collection of
(300–)550–2500(–2900) m a.s.l. In Belize it is found seed and specimens was carried out by the Oxford
between 300–760 m a.s.l. It is a major constituent of Forestry Institute (OFI), as well as by other organi-
more or less open to closed-canopy pine and pine- zations, throughout its entire range. A major limita-
oak forests in climatic zones which receive at least tion to introductions on a large scale is the limited
1000 mm of rainfall per annum, and up to 2500– availability of seeds, both from natural stands and
3000 mm in some places. The dry season is usually from so-called seed orchards. It has been planted as
long, lasting from November to May, so that at lower a forestry tree in Africa, India, South America and
to middle altitudes fires are an integral phenomenon Australia (Queensland). It is not known to be used
in the ecosystem, the frequency of which has been as an amenity tree.
however greatly accelerated by humans. Here, open
 Pinus teocote Schiede ex Schltdl. & Cham., Linnaea
Distribution
5: 76. 1830. Type: Mexico: Veracruz, Pico de
Orizaba, (Volcan Citlaltepetl), C. J. W. Schiede & Mexico: Chihuahua, Coahuila, Nuevo León, Tama-
F. Deppe s.n. (holotype HAL). ulipas, Sinaloa, Durango, Zacatecas, San Luís Potosí,
Nayarit, Aguascalientes, Jalisco, Guanajuato, Queré-
taro, Hidalgo, Michoacán, México, Distrito Federal,
Etymology
Tlaxcala, Puebla, Veracruz, Guerrero, Oaxaca and
Teocote was apparently one of the Aztec names used Chiapas.
for pines in Mexico. TDWG codes: 79 MXC-DF MXC-ME MXC-PU MXC-
TL MXE-AG MXE-CO MXE-CU MXE-DU MXE-GU
772 MXE-HI MXE-NL MXE-QU MXE-SL MXE-TA MXE-
Vernacular names
ZA MXG-VC MXN-SI MXS-GR MXS-JA MXS-MI MXS-
Aztec pine, Teocote pine; pino chino, pino colorado, NA MXS-OA MXT-CI
pino real, pino rosillo (Spanish)
Ecology
Description
Pinus teocote occurs in various habitats, most com-
Trees to 20–25 m tall, d.b.h. to 75 cm; trunk mono­ monly in rather open forest or woodland associated
podial, sometimes forked, straight. Bark thick, with Quercus spp., on relatively dry sites with shal-
rough and scaly, forming longitudinal plates divided low soils. Its altitudinal range is (1000–)1500–3000
by deep, wide fissures, dark greyish brown. Branches (–3300) m a.s.l., a few collections are recorded from
spreading horizontally or curved down; branches of 700–800 m but these may be based on incorrect esti-
higher orders slightly pendulous, forming a pyrami- mates. This species often occupies dry ridges, some-
dal to rounded crown. Shoots smooth, orange-brown. times on calcareous outcrops. Annual precipitation
Cataphylls subulate, curved at apex, with ciliate is usually moderate, from 500–1000 mm, but much
margins, brown. Vegetative buds ovoid-oblong; ter- higher in parts of the central highlands of Chiapas,
minal bud 10–15 mm long; lateral buds ovoid-acute, where it is growing in broad-leaved forest dominat-
smaller, all not resinous. Fascicle sheaths initially up ed by Liquidambar and Styrax. Pinus teocote occurs
to 20 mm long, in mature fascicles reduced to ca. with a number of other pines in various parts of its
10 mm. Leaves in fascicles of 3(2–5), persisting 2–3 range, the most common of which are from N to S:
years, straight or slightly curved, rigid, (7–)10–15 P. arizonica, P. engelmannii, P. durangensis, P. leio­
(–18) cm long, 1–1.4 mm wide, with serrulate mar- phylla, P. montezumae, P. oocarpa, and P. patula.
gins, acute-pungent, light green. Stomata on all faces
of leaves. Pollen cones ovoid-oblong to cylindrical,
Conservation
1–1.8 cm long, ca. 5 mm wide, yellowish green. Seed
cones commonly opposite, sometimes 1–3, on short, IUCN: LC
stout, curved peduncles falling with cones. Mature
cones ovoid to ovoid-oblong when closed, slightly
Uses
asymmetrical, with a broad, flattened but oblique
base when opened, (3–)4–6(–7) × 2.5–5 cm when Aztec pine is of importance as a timber tree. Appar-
open, often persistent. Seed scales thick woody, ob- ently due to its moderate size especially in the south-
long, straight or recurved. Apophysis slightly raised, ern part of its range, resin production may locally
in some cones more or less flat, transversely keeled, count as the more important mode of exploitation.
apical margin angular or crenate, light brown, in The wood is used as sawn timber, mainly for rail-
some cones with radial marks. Umbo dorsal, flat to way sleepers, coarse construction work, containers
blunt-pyramidal, with a minute, deciduous prickle. and crates, particleboard, and also for pulp. Resin
Seeds 3–5 mm long, dark grey-brown; 12–18 × 6–8 tapping is still an important industry, but it will
mm, translucent, yellowish with a dark tinge. ­fluctuate much with market prices for naval stores.
In horticulture this species is hardly known, despite dispersal, ovoid-conical when closed, 4–6(–7) cm
its wide range geographically and altitudinally, indi- long, opening to (broadly) ovoid, 3–4.5 cm wide.
cating hardiness of at least some provenances. Har- Seed scales woody, rigid, oblong; apophyses nearly
diness requirements, of course, make the implicit flat to slightly raised, transversely keeled, rhombic
assumption that only people living in cool or cold or with a rounded upper margin, light brown, more
winter climates want to (or should) grow trees. In or less lustrous; umbo dorsal, small, unarmed. Seeds
addition, notions written in conifer handbooks like obovoid, slightly flattened, 5–7 mm long, grey-
“least attractive of Mexican pines” first of all fail to brown; wing oblong, 10–15 mm long, pale brown
acknowledge that taste is a personal thing and sec- with dark stripes.
ond do no good to promote its cultivation, which
may well be of interest to dendrologists. 773
Distribution
Japan: Honshu, Kyushu, Shikoku; South Korea (near
Pinus thunbergii Parl., in Candolle, Prodr. 16 (2): the coast).
388. 1868. Pinus thunbergiana Franco, Anais Inst. TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-SK
Super. Agron. (Lisboa) 16: 130. 1949. Type not
designated.
Ecology
This is a species of pine growing at low to middle
Etymology
elevations (up to ca. 1000 m above sea level) in the
This species was named after Carl Peter Thunberg coastal hills and mountains of the islands of Japan
(1743–1828), who published an early Flora of Japan. and South Korea, where the climate is warm tem-
perate (with little or no frost) and moist. These re-
gions would have had a predominantly deciduous
Vernacular names
angiosperm forest cover, with conifers mixed in
Japanese black pine; kuro-matsu (Japanese); haesong especially on poor, water-logged soils and on dry
(Korean) slopes and mountain ridges. Pinus thunbergii would
have occupied these habitats as well as those in close
proximity to the sea coast. Extensive cultivation has
Description
removed the natural vegetation in most areas, but as
Trees to 40 m tall; trunk to 2 m d.b.h., often forked a pioneer species P. thunbergii has been able to hold
in crown. Bark on trunk thick and breaking into its own; it has been much planted in afforestation
scaly ridges separated by longitudinal, deep fissures, schemes from where it could spread in adjacent un-
grey with a purplish hue, nearly black in the fissures. cultivated areas. Its tolerance of salty winds makes it
Branches spreading and ascending, sometimes a species that grows well on the sea coasts of Japan,
heavy, forming broadly conical to domed crowns. both naturally and when planted; naturally its trunk
Foliage branches stout, rough with pulvini from becomes bent and the crown flattened under severe
fallen leaf fascicles, glabrous, new shoots yellowish exposure.
green, becoming light brown or yellowish brown
to grey. Buds ovoid-oblong to ellipsoid-cylindrical,
Conservation
acuminate, 12–20 mm long, non-resinous or resin-
ous; cataphylls light brown, thin, with papery, grey- IUCN: LC
white to silver-white fringes. Leaves in fascicles of
2, held by a persistent, 10–12 mm long basal sheath,
Uses
remaining 2–3 years on branchlets, straight, rigid,
(6–)7–12 cm long, usually twisted, 1–2 mm wide, The wood of this pine is similar to that of the Black
dark green; margins minutely serrate; apex acute pine (Pinus nigra) and is used for general construc-
or pungent; stomata in fine lines on all faces. Pollen tion, poles, railway sleepers, fences, pallets and
ovoid-conical to short cylindrical, 1–1.5 cm long, 0.5 crates, flooring, fibreboard, and wood pulp. Japa-
mm wide, yellow; Seed cones solitary or in whorls nese black pine is mostly used as a windbreak tree
of 2–3 on short peduncles, falling shortly after seed and to stabilize sand dunes in coastal areas near
 ­ rbanization. It is also widely planted as an orna-
u Seed scales thick woody, rigid, cuneate, with 2 deep
mental tree in Japan and Korea (somewhat less com- seed cavities adaxially; apophyses strongly devel-
monly in other countries) and some cultivars have oped, markedly raised, multi-­angular or more or
been selected especially in Japan to suit traditional less rhombic, sharply transversely keeled or with
Japanese gardening. It is also being used in bonsai 4–5 convergent ridges, lustrous light or red-brown;
growing. In the USA this species was widely planted umbo dorsal and central, pyramidal or slightly
for afforestation in coastal areas of New England, curved, with an obtuse, hard apex. Seeds large,
until susceptibility to pests and diseases put an end obovoid, slightly flattened, 17–24 mm long, 10–14
to these schemes and forced the species back to ar- mm wide, light brown or darker mottled; wing re-
boreta and parks, where solitary trees are usually duced, up to 10 mm long, surrounding the seed like
774 safe. Minor uses in Korea are of the needles in pastry a partial ring, deciduous and sometimes remaining
and in (medicinal) soft drinks. attached to the seed scale.

Distribution
Pinus torreyana Parry ex Carrière, Traité Gén.
Conif.: 326. 1855. USA: S California (San Diego and Santa Barbara
Co.).
TDWG codes: 76 CAL
Etymology
This species was named by W. E. Parry, who failed to
Ecology
validly publish it, after the American botanist John
Torrey (1796–1873). Pinus torreyana is a relict species now confined to
littoral habitat on the coast (up to 1.6 km inland) and
on a small island off the coast of southern Califor-
Vernacular names
nia. It grows from immediately above the high tide
Torrey pine, Soledad pine, Del Mar pine mark to about 180 m a.s.l. on rocky or sandy slopes.
On these sites it seems dependent on the daily fog
that comes in from the ocean in the afternoon, miti-
Description
gating the heat of the sun and the resulting excessive
Trees to 15(–23) m tall; trunk to 1 m d.b.h., mostly evapo-transpiration. It grows with a sparse chapar-
curved or crooked in habitat. Bark rough and scaly, ral and few other trees; in ravines sometimes ac-
deeply fissured on trunk, reddish brown turning to companied by a few oaks (Quercus sp.) and Arbutus
purplish grey. Branches spreading and assurgent or menziesii.
ascending, irregular, forming an open or flattened
crown. Foliage branches stout, ca. 10 mm thick (20
Uses
mm thick on coning shoots), new shoots glabrous,
greenish, becoming purplish brown to nearly black. Torrey pine is not used as a timber tree; at present
Buds ovoid-conical, 20–30 mm long, without resin; the two disjunct populations are protected by law. It
cataphylls light brown with white fringed margins. is in cultivation in California in gardens and some
Leaves in fascicles of 5, rarely 4 or 6, held in up to arboreta, but rare elsewhere. In the better growing
20 mm long but later shortening, persistent basal conditions of gardens it can grow to a large tree; a
sheaths, persisting 3–4 years, spreading, rigid, 15– specimen in New Zealand was 45 m tall with a d.b.h.
25 cm long, straight or curved, slightly twisted, ca. of 1.5 m in 1982 (Grimshaw & Bayton, 2009: 626).
2 mm wide, greyish green; margins minutely serru- Although its conservation seems more or less cared
late; apex abruptly acute; stomata in lines on all fac- fore at present, growing this species more widely as
es. Pollen cones ovoid-cylindrical, 2–3 cm long, 8–10 an ex situ backup is to be recommended; it is also
mm wide, yellow. Seed cones solitary or rarely in an interesting species to grow and requires a mild
pairs on stout, 3–4 cm long peduncles, maturing in climate with warm, sunny summers and (near) ab-
3 years, persisting to 5 years, massive, 10–16 cm long, sence of frost in winter.
broadly ovoid to ovoid-conical, opening slowly and
only partially to 8–17 cm wide, strongly ­resinous. 2 subspecies are recognized:
 775

3 2

Pl ate 3 4. Pinus torreyana subsp. torreyana. 1. Habit of tree. 2. Leaves. 3. Seed cone. 4. Seed.
 Pinus torreyana Parry ex Carrière subsp. torreyana. TDWG codes: 76 CAL
Type: USA: California, San Diego Co., Torrey Pines
State Park, J. R. Haller 10450 (neotype UCSB). Pl. 34
Conservation
This tiny population of ca. 1000 mature trees cov-
Description
ering less than 100 ha in two subpopulations on a
Trees with erect trunks commonly 15, sometimes near-shore island is at high risk from stochastic
more than 20 m tall. Seed cones ovoid when closed, events such as fire, pest infestation, or disease, all
less than 12 cm wide. Seeds more or less evenly of which could wipe out this subspecies easily. The
brown, not or only lightly mottled. trees enjoy legal protection, but that is not an effec-
776 tive buffer against these risks.
IUCN: EN (D)
Distribution
USA: S California (San Diego Co., on the coast N of
San Diego). Pinus tropicalis Morelet, Rev. Hort. Côte d’Or 1:
TDWG codes: 76 CAL 106. 1851. Type: Cuba: Isla de la Juventud [Isla de
Pinos], La Cañada, N. L. Britton & E. G. Britton
14416 (neotype NY).
Conservation
The small population on the mainland that constitutes
Etymology
this subspecies is in part (southern subpopulation)
legally protected in the Torrey Pines State Park. How- The species epithet means that this pine occurs in
ever, the small overall size, fewer than 3500 mature the tropics.
trees covering ca. 320 ha in two subpopulations, and
close proximity to major urban development, put the
Vernacular names
subspecies highly at risk of destructive events such as
fires, pest epidemics and diseases. Trees outside the Tropical pine
reserve are often not protected from development;
they are sometimes incorporated in urban landscap-
Description
ing and sometimes felled (personal obs. 1992).
IUCN: EN (C2b) Trees to 30 m tall, d.b.h. to 1(?) m; trunk monopo-
dial, erect. Bark thick, rough, scaly, breaking into ir-
Pinus torreyana Parry ex Carrière subsp. insularis regular polygon plates, divided by deep longitudinal
J. R. Haller, Syst. Bot. 11 (1): 45. 1986. Pinus tor­ fissures on lower part of the trunk, reddish brown,
reyana Parry ex Carrière var. insularis (J. R. Haller) weathering grey. Branches spreading to ascending,
Silba, Phytologia 68: 64. 1990. Type: USA: Cali- forming an irregular, open crown. Shoots uni-nodal,
fornia, Santa Barbara Co., Santa Rosa Island, thick, very rough, lustrous orange-brown in 1st and
J. R. Haller 10448 (neotype UCSB). 2nd year, then grey. Cataphylls subulate, recurved,
scarious, brown. Vegetative buds ovoid-oblong,
acute; terminal bud 15–25 mm long; lateral buds
Description
smaller, not resinous. Fascicle sheaths initially ca. 20
Stunted trees commonly less than 10 m tall, with a mm long, persistent but reduced to ca. 10 mm on
flat crown (due to severe growing conditions). Seed mature leaf fascicles. Leaves in fascicles of 2 (rarely
cones broad when closed, usually over 12 cm wide, 3), persisting 2 years, very uniform, straight and rig-
with strong, nearly straight umbos. Seeds with dark id, (15–)20–30 cm long, 1.5 mm wide, with serrulate
spots. margins, acute, light or yellowish green. Stomata on
all faces of leaves. Pollen cones ovoid-oblong to cy-
lindrical, 2–3 cm × 5 mm, pink to yellowish. Seed
Distribution
cones subterminal, on short, thick peduncles, soli-
USA: S California (Santa Barbara Co., Santa Rosa tary, in pairs or whorls up to 6, remaining erect or
Island). spreading obliquely. Mature cones narrowly ovoid to
ovoid-attenuate when closed, ovoid with a flattened Pinus uncinata Ramond ex DC., in Lamarck &
base when opened, then 5–8 × 4–5.5 cm, persistent Candolle, Fl. française, ed. 3, 3: 726. 1805. Type:
for some years, but falling with peduncle attached. Europe: [locality not mentioned], K. I. Christensen
Seed scales oblong, straight or strongly recurved. F6, 5/12 July 1980 (neotype G).
Apophysis flat or slightly raised, transversely keeled,
rhombic to pentagonal in outline, radially striated, Pinus uncinata Ramond ex DC var. rostrata An­toine,
light brown or reddish brown. Umbo dorsal, flat or Conif.: 12. 1840; Pinus mugo Turra var. rostrata (An-
slightly raised, without a prickle. Seeds 5 × 4 mm, toine) Hoopes, Ber. Deutsch. Bot. Ges. 58a: 20. 1941;
light grey-brown; wings 12–15 × 5–6 mm, yellowish Pinus mugo Turra subsp. rostrata (Antoine) E. Mur-
with a grey or black tinge. Seedling: Suppressed ter- ray, Kalmia 13: 23. 1983.
minal growth and a thick radicle produce a ‘grass Pinus uncinata Ramond ex DC var. ancestralis 777
stage’ as an adaptation to ground fires. Leaves are Businský, Acta Pruhon. 88: 7. 2008.
normal but less straight.
Etymology
Distribution
The species epithet comes from the Latin uncinus =
Caribbean: W Cuba (Pinar del Rio, Isla de la Juven- barb, referring to the hooked apophyses on the cone
tud [Isla de Pinos]). scales.
TDWG codes: 81 CUB
Vernacular names
Ecology
Mountain pine; pino negro (Spanish); pin à crochets
Pinus tropicalis is a lowland pine, occurring on the (French); Aufrechte Bergföhre, Hakenkiefer (Ger-
coastal plains and low foothills between 1–150(–300) man).
m a.s.l., on nutrient-poor sandy or gravelly alluvial
soils which are dry due to rapid drainage. The cli-
Description
mate is tropical, with annual precipitation of around
1200 mm and a prolonged dry season. It is in part Trees to 20–25 m tall (often smaller); trunk to 50 cm
sympatric with P. caribaea var. caribaea, which has d.b.h. Bark breaking into small, rectangular plates
a greater altitudinal range. Pine savanna’s are open, and sometimes curling scales on larger stems, dark
grass-dominated lowland areas which burn fre- brown. Foliage branches stout, rough with decur-
quently; P. tropicalis has an advantage over P. carib­ rent pulvini and small cataphylls, glabrous, at first
aea through its ‘grass stage’ by which the seedling orange brown, later reddish brown. Buds ovoid-
can survive successive fires. Thus it becomes fre- cylindrical, acute, strongly resinous, 5–10 mm long.
quently the only pine in this vegetation type. Leaves in fascicles of 2, held in 10–18 mm long, later
much shorter, basal fascicle sheaths, densely set on
branches, directed forward, persisting 4–8 years, (3–)
Conservation
5–6 cm long, curved and rigid, twisted, 1.5–1.8 mm
IUCN: LC wide, dark green; margins minutely serrulate; apex
acute or pungent; stomata in fine lines on all sides.
Pollen cones short cylindrical, 10–15 mm long, 5 mm
Uses
wide, yellow. Seed cones solitary or in whorls of 2–3,
Pinus tropicalis is an important regional source of sessile or on very short peduncles below shoot apex,
timber mainly used by local sawmills. Its wood is patent or more or less reflexed, falling at maturity or
dense and durable, but resinous. Despite extensive persisting up to 4 years, mostly oblique when closed,
exploitation, it is still common. Regeneration seems 3–6.5 cm long, 3–5(–6) cm wide when opened, with
to be good in many areas especially where it is pro- an asymmetrical, flattened base. Seed scales thin
tected from grazing pressures. It is also used in plan- woody, rigid, oblong, spreading wide, dark brown
tation forestry, mainly in Cuba, but also to a limited or blackish brown. Apophyses prominently raised,
extent elsewhere. transversely keeled, on outer, sun-exposed side of
cone much larger and recurved or longitudinally
 ridged (‘ barbed’ or ‘hooked’), lustrous yellowish Calluna vulgaris, Eriophorum vaginatum, and some
or light brown. Umbo dorsal, conical or obliquely oligotraphent species of Sphagnum like S. recurvum
pyramidal, excentric, armed with a minute prickle. and S. magellanicum, most of these with a high pres-
Seeds small, 3–4 mm long; wing articulate, 10–13 ence in the vegetation. Pinus uncinata occurs natu-
mm long. rally at altitudes of between 600 and 1600 m a.s.l. In
the Alps and Pyrenees it grows on moist slopes up to
the tree line; in the Alps it is confined to the western
Taxonomic notes
and northwestern ranges that are much wetter than
This species has often been considered to be a va- the central and southern Alps. Here it sometimes
riety or subspecies of Pinus mugo and has been grows with Picea abies or Pinus sylvestris; in the Pyr-
778 classified under all its synonyms and possible com- enees the spruce is naturally absent.
binations at these ranks (see Farjon, 1998, [2001] for
full synonymy under P. mugo subsp. uncinata). The
Conservation
principal distinction with P. mugo is the tree habit of
P. uncinata and not the oblique cones with strongly IUCN: LC
developed apophyses on one side, as these are found
in some of the shrubby mountain pines as well
Uses
(see P. mugo subsp. rotundata on p. 718 and Chris-
tensen, 1987). I have maintained this taxon in this Mountain pine is not commercially important; its
Handbook as a distinct species, as I did in the sec- main value is ecological and it has some significance
ond edition of my book Pines (Farjon, 2005b). The as an ornamental tree. The wood is used as fuel and
tree habit is not attributable to ecological factors, as is suitable for light construction, but as this species
it is constant through the entire altitudinal range grows slowly and remains small the yield is limited.
in which this pine occurs and is maintained when Its only commercial use is located in the Pyrenees,
seeds are grown up under controlled conditions, as where fairly extensive stands exist. Some of the wood
is the case with the shrubby habit of P. mugo and its is excellent for special uses like turnery, woodware
subspecies or varieties. Both taxa can and do form and musical instruments due to its relative density
hybrids in nature with P. sylvestris and probably with and hardness. It is uncommon in cultivation and
each other. One would hope that taxonomists could restricted to local parks and further afield to some
finally arrive at a consensus, but that hope seems botanic gardens and arboreta.
idle and the 200-year old tangle of names and their
synonyms is likely to be expanded with yet more
new names and combinations applied to this group Pinus virginiana Mill., Gard. Dict., ed. 8: Pinus No.
of European pines. 9. 1768. Type not designated. Fig. 251

Distribution Etymology
Europe: NE Spain, Pyrenees, Auvergne Mts. (pos- This species name indicates its native origin, Virgin-
sibly introduced), Alps, Jura, Vosges, Böhmerwald, ia, with which much of the eastern USA was desig-
Erzgebirge. nated in colonial times.
TDWG codes: 11 AUT-AU CZE-CZ GER SWI 12 FRA-
FR SPA-AN SPA-SP
Vernacular names
Virginia pine, Scrub pine
Ecology
This is a pine of wet moors in W European mountains
Description
and typical for the association Pinetum uncinatae
Kastn., Flössn. & Uhlig (= Vaccinio uliginosi-Muge- Trees to 15–18 m tall, rarely to 25 m; trunk to 50 cm
tum Oberdorfer). This association has only Pinus d.b.h., straight or contorted, forked, or shrubby. Bark
uncinata as a characteristic species, but it is usually thin and scaly, irregularly fissured and ridged on
accompanied by various species of ­Vaccinium, by lower part of larger trunks, grey-brown, becoming
reddish higher up in the tree. Branches numerous, it can attain taller tree stature in mixture with other
irregularly spreading and ascending, forming an ir- trees in a forest environment. In such woods it is a
regular, often dense, rounded or flat-topped crown. minor component accompanying Quercus spp. and
Foliage branches slender, rough with pulvini from sometimes other species of pine, e.g. P. echinata,
fallen leaf fascicles, new shoots glabrous, purplish P. rigida and P. taeda, or it is part of a mixture of
brown, often glaucous, becoming red-brown to grey. oaks and pines. Other conifers locally growing with
Buds short and broad, ovoid, 5–10 mm long, shortly P. virginiana are Tsuga canadensis and Thuja occi­
acute, resinous or not; cataphylls red-brown, with dentalis. In the so-called pine barrens in the NE of
whitish fringed margins. Leaves in fascicles of 2, held its range sometimes lichens (Cladina, Cladonia) and
by short, persistent sheaths, remaining 3–4 years a few oak shrubs (Quercus ilicifolia) provide the only
on branches, spreading, rigid, 3–8 cm long, often undergrowth. 779
strongly twisted, 1–1.5 mm wide, light or dark green
or yellowish green; margins minutely serrulate; apex Conservation
narrowly acute; stomata in inconspicuous lines on
IUCN: LC
all surfaces. Pollen cones ellipsoid-cylindric, 10–15
mm long, red-brown turning yellow at anthesis. Seed
Uses
cones solitary or in pairs, nearly sessile or on 1 cm
long, tenacious peduncles, persistent but opening Foresters dismissed this species as ‘weedy’ in the
soon, spreading out or curved back, 3.5–7 cm long, past, but it has the capacity to grow upright to a
slightly oblique to symmetrical, conical when closed, moderate size, especially when planted on aban-
ovoid when open. Seed scales woody, rigid, oblong, doned farmland and former coal strip mines. It
dull red-brown with a purple-red sealing band; apo- provides rough lumber and pulpwood on these sites
physes prominently raised or slightly thickened, especially in the SE of its range. It is also important
transversely keeled, light brown; umbo dorsal and as a Christmas tree and huge quantities are planted
central, pyramidal, armed with a slender, strong, to and sold each year. It finds little appreciation in hor-
5 mm long, straight or curved prickle. Seeds obovoid, ticulture, but the dense branching of some shrubby
slightly flattened, 4–6(–7) mm long, pale brown with specimens would be ideal in garden landscaping.
darker spots; wing 16–20 mm long, narrow. Dwarf forms (cultivars) from witches brooms have
been produced by growers in the USA and can be
used to advantage in rock gardens.
Distribution
E USA: from New York State in the north to N Ala-
bama and Mississippi in the south-west and South Pinus wallichiana A. B. Jacks., Bull. Misc. Inform.,
Carolina in the south of its range. Kew 1938: 85. 1938.
TDWG codes: 75 INI NWJ NWY OHI PEN WVA 78
ALA DEL GEO KTY MRY MSI NCA SCA TEN VRG
Etymology
This species was named in honour of Nathaniel Wal-
Ecology
lich (1786–1854), a Danish botanist who amassed the
Pinus virginiana is a species of the Piedmont and famous Wallich Herbarium for the East India Com-
lower slopes of the Appalachian Mountain system, pany, now kept at K.
growing on the sea coast in the north, but only in
the interior and at higher altitudes in the south of
Vernacular names
its range, up to 650 m a.s.l. The climate is humid and
cool in most of its range and snowfall can be abun- Himalayan white pine, Blue pine, Bhutan pine; kail
dant at least in the northern parts. This species is (Hindi); lamshing (Kumaon, Bhutan)
naturally restricted to poorer soils and avoids cal-
careous substrates. It is a pioneer that now invades
Description
large tracts of abandoned farmland, invigorating its
reputation as a weedy species. On these and other Trees to 50–60(–70) m tall; trunk to 2 m d.b.h., bole
marginal or disturbed sites it is a shrubby tree, but straight and columnar. Bark becoming fissured and
 breaking into small scales that flake off in small chips, broad-leaved trees, e.g. species of the genera Quer­
dark greyish brown. Branches whorled, spreading cus, Acer and Ilex. In the western Himalayas, where
wide and curved downward, those of higher orders conditions are drier, P. wallichiana forms mixed for-
drooping but with upturned ends, forming a broad- ests with Cedrus deodara. Other conifers with which
ly pyramidal crown. Foliage branches glabrous, new it may be associated are Pinus roxburghii, Abies
shoots at first glaucous green, becoming pale green spectabilis, or A. densa and Tsuga dumosa in the wet-
and then grey-brown. Buds ovoid-conical; termi- ter eastern part of its range.
nal bud more or less conical, 10–15 mm long; lateral
buds smaller and more ovoid, all slightly resinous.
Uses
Leaves in fascicles of 5, held in deciduous, basal fas-
780 cicle sheaths of orange-brown flimsy scales, persist- Bhutan pine is an important timber tree in many
ing 2–3 years, slender and flexible, (6–)10–18(–20) parts of the Himalaya. It is of similar timber prop-
cm long, drooping or sometimes pendulous, often erties and quality to P. strobus and P. monticola in
with a sharp bend near base but otherwise straight, North America, with tall, straight trees producing
1 mm wide; margins minutely serrulate, (dark) straight grained wood of good strength. It is used for
green abaxially, glaucous white adaxially; stomata construction, carpentry and joinery, wall panelling,
in fine lines on the two adaxial faces. Pollen cones veneers, furniture, fences and gates, crates and box-
in small clusters at base of new shoots, with 8–12 es, and railway sleepers after treatment with preserv-
basal perular scales remaining and partly cover- atives. In India (Himachal Pradesh) resin tapping
ing the short cylindrical, yellow cones. Seed cones is an important use to obtain naval stores. A sweet
solitary or in whorls of 2–6, initially erect on stout, liquid known as honey dew is secreted by aphids
2.5–4 cm long peduncles, becoming pendulous from the leaves and collected by local people of the
when growing, cylindrical, usually slightly curved, mountain forests for consumption. Bhutan pine was
(10–)15–30 cm long, 3–4 cm wide when closed and introduced to England in 1823 and, unlike several
5–8(–9) cm wide when mature and opened, usually other species of Pinus subsection Strobi, it turned
resinous, soon falling with peduncle after seeds have out to be relatively immune to infections with blister
been released. Seed scales cuneate-oblong, widest rust (Cronartium ribicola; Basidiomycota) as well as
just ­below the apophysis, thin woody, only slightly to atmospheric pollution. In forestry it is also used
flexible, with two seed cavities near adaxial base. in plantations and several hybrids with related spe-
­Apophyses more or less rhombic, slightly raised, cies have been established with timber production
with grooves converging to the terminal, small, in mind (e.g. the cross between P. strobus and P. wal­
incurved, obtuse-triangular umbo, lustrous light lichiana = P. ×schwerinii Fitschen). Bhutan pine is a
brown with darker umbo. Seeds obliquely obovoid, widely used amenity tree and a number of cultivars
(3–)5–9 mm long, 3.5–5 mm wide, yellowish brown have been selected and are in the trade.
or brown; wing (10–)20–35 mm long, (5–)8–10 mm
wide, grey-brown. 2 varieties are recognized:

Pinus wallichiana A. B. Jacks. var. wallichiana.

Distribution
Type: Illustration in Lambert, Descr. Pinus 2, t. 3
Himalaya: from Afghanistan (Hindu Kush) to NE “Pinus excelsa”. 1824 (lectotype designated here).
India, SE Xizang [Tibet] and NW Yunnan. Fig. 252
TDWG codes: 34 AFG 36 CHC-YN CHT 40 EHM-AP
EHM-BH NEP PAK WHM-HP WHM-JK WHM-UT Pinus wallichiana A. B. Jacks. var. manangensis
H. Ohba & M. Suzuki, [Himalayan Pl. 1] Univ. Mus.
Univ. Tokyo Bull. 31: 350. 1988.
Ecology
Pinus wallichiana grows in the Himalayas in the val-
Description
leys and foothills, to a maximum altitude of 2700 m,
but in Bhutan it reaches 3400 m a.s.l. Sometimes it Leaves 10–20 cm long. Seed cones 15–30 cm long.
forms pure stands or forests, in other places it ap- Seeds 4–9 mm long, with a 15–35 mm long and 8–10
pears as an important forest component mixed with mm wide wing.
 Distribution Vernacular names
Himalaya: from Afghanistan (Hindu Kush) to Bhu- mao zhi wu zhen song (Chinese)
tan; China: SE Xizang [Tibet].
TDWG codes: 34 AFG 36 CHT 40 EHM-BH EHM-DJ
Description
EHM-SI NEP PAK WHM-HP WHM-JK WHM-UT
Trees to 20 m tall; trunk to 60 cm d.b.h. Bark on
young trees and branches smooth, thin, becoming
Conservation
scaly and flaking, brown, dark brown or grey-brown
IUCN: LC on trunks of larger trees. Branches spreading wide,
forming broad, umbrella-shaped or irregular, flat- 781
Pinus wallichiana A. B. Jacks. var. parva K. C. Sahni, topped crowns. Foliage branches slender; young
Indian J. Forest. 12 (1): 40. 1989. Type: India: Aru- shoots red-brown, initially pubescent, glabrous
nachal Pradesh, Kameng District, Tawang, R. N. Lo­ or with remnants of pubescence in grooves in the
ganey 6 (holotype DD). second to third year, turning greyish brown. Buds
ovoid to cylindrical, not resinous; cataphylls brown.
Leaves in fascicles of 5, held by deciduous sheaths
Description
of flimsy, brown scales, spreading or curved slight-
Leaves 6–11(–13) cm long. Seed cones ca. 10 cm long. ly towards shoot, 2.5–6 cm long, pliant, straight or
Seeds consequenly small, ca. 3 mm long, with a curved, 1–1.5 mm wide, green, with stomatal lines on
10 mm long and 5 mm wide wing. the two adaxial surfaces; margins minutely serrulate.
Pollen cones in small clusters, short cylindrical. Seed
cones variable in size and shape, from short ovoid to
Taxonomic notes
long cylindrical, initially erect on stout peduncles,
This apparently high altitude form from NE India in becoming curved down to pendulous, 4–10(–15)
an area also claimed by China as lying in Xizang [Ti- cm long, solitary or with 2–3 together. Seed scales
bet], see Flora of China 4: 24 (1999), was described soft woody, more or less flexible at base, cuneate to
from a type specimen only. The area remains inac- oblong; apophyses rhombic to oblong (at base and
cessible due to the disputed international border. apex of cone), curved or more or less straight, not
recurved or more commonly recurved near cone
base, ripening to yellowish brown or dark brown,
Distribution
weathering grey-brown; apex thin and straight or
NE India: Arunachal Pradesh (Assam, Kameng Dis- slightly incurved, rarely upcurved; umbo terminal,
trict). small and sunken or obtuse. Seeds obovoid or ellip-
TDWG codes: 40 EHM-AP soid, 8–10 mm long, ca. 6 mm wide, usually with a
well developed wing ca. 16 mm long and 7 mm wide.
Conservation
Taxonomic notes
IUCN: LC
Pinus wangii has been accepted as a distinct species
in Flora of China 4 (1999) and in the World Check-
Pinus wangii Hu & W. C. Cheng, Bull. Fan Mem. list and Bibliography of Conifers (Farjon, 1998,
Inst. Biol., ser. 2, 1 (2): 191. 1948. Type: China: [2001]) but was only tentatively accepted by Farjon
Yunnan, Xichou Xian, Fadou, C. W. Wang 85830 (2005b) in the description of P. fenzeliana Hand.-
(holotype KUN). Mazz. A complex of closely related taxa (among
which also feature P. kwangtungensis and P. dalaten­
sis) occurs in SW China and Vietnam. The Czech
Etymology
botanist R. Businský (2008) has treated these taxa
The species epithet commemorates the Chinese bot- in a narrowly circumscribed concept of species and
anist C. W. Wang, who collected the type specimen subspecies, recognizing more species than most.
in SE Yunnan, China. Hu & Cheng (op. cit.) compared their new species
 to P. parviflora of Japan, but we now consider that
Description
species more distantly related. The morphology is
highly variable in some of these species and may not Trees to 30 m tall, rarely shrubs; trunk to 1 m d.b.h.,
be sufficiently clear to resolve the taxonomy; addi- in trees monopodial, sometimes forked in crown.
tional DNA analysis may be needed, but could only Bark scaly, longitudinally fissured, breaking into
help to resolve this species problem if based on suffi- irregular plates, greyish brown weathering grey,
ciently wide sampling. Ideally, clades resolved from flaking in small or large chips. Branches spreading
DNA data should correlate with groupings based on or curving down, forming a domed or flat-topped
distinct morphological character states, and only crown. Foliage branches stout, becoming rough with
groups that are congruent on both criteria should be pulvini after shedding leaves, glabrous, new shoots
782 given species names. thick, often vigorous, reddish brown, becoming
brown or grey-brown. Buds ovoid-oblong, acute;
terminal bud to 25 mm long; lateral buds smaller,
Distribution
not resinous; cataphylls appressed, red-brown.
China: SE Yunnan (Malipo, Xichou); Vietnam (Mai Leaves in fascicles of (2–)3, more or less remote,
Chou) (?) [The taxonomic status in Vietnam re- persisting 2–3 years, spreading, held in a 10–15 mm
mains unclear, see Hiep et al., 2004)]. long, persistent basal sheath, 7–20(–30) cm long,
TDWG codes: 36 CHC-YN 41 VIE straight and slender, pliant, only very long leaves
drooping, 1–1.2 mm wide, often slightly twisted,
light green; margins minutely serrulate; apex acute
Ecology
or acuminate; stomata in fine lines on all surfaces.
Pinus wangii is probably restricted to the ‘limestone Pollen cones short cylindrical, ca. 2 cm long, yellow.
country’ of S China, where it occurs on steep slopes Seed cones in pairs or whorls of up to 5, short pe-
and limestone ridges, associated with Quercus varia­ dunculate or nearly sessile, usually persisting a few
bilis and other, mostly small-leaved, evergreen trees years but opening soon, ovoid-conical when closed,
and shrubs. Its altitudinal range there is from 500 to near-symmetrical, 5–10(–11) cm long and 3.5–4.5 cm
1800 m a.s.l. wide, opening to 5–7 cm wide. Seed scales woody,
rigid, oblong to obovate, except basal scales spread-
ing wide. Apophyses raised, variously shaped, trans-
Conservation
versely keeled, ripening from green to light brown or
IUCN: EN (B1+2b+d) chestnut-brown; umbo nearly flat or raised, armed
with a short or minute prickle. Seeds ovoid, 4–5 mm
long, brown; wing 12–15 mm long, 5 mm wide, light
Uses
brown or grey-brown.
Apparently used locally as a timber tree, no specifics
are known. This species is rare in cultivation; there
Taxonomic notes
are a few, mostly young, trees in botanic gardens in
China, the UK and the USA. This species resembles and hybridizes with Pinus
tabuliformis, forming the hybrid taxon Pinus ×den­
sata Mast. Some authors have considered it a subspe-
Pinus yunnanensis Franch., J. Bot. (Morot) 13 (8): cies of P. kesiya (or a variety of P. insularis = P. kesiya
253. 1899. var. langbianensis). It would require detailed genetic
(DNA) analysis to establish the true relationships of
these taxa, and it seems more appropriate to retain
Etymology
this taxon at species rank at present. A form with
The species epithet denotes Yunnan, China, from very long, drooping needles up to 30 cm long was
where it was first described. named Pinus yunnanensis var. tenuifolia and re-
ported to occur in Guangxi and Guizhou, China, at
400–1200 m a.s.l. It is recognized as distinct in Flora
Vernacular names
of China 4: 15 (1999), but the characters stated to dif-
Yunnan pine; yun nan song (Chinese) ferentiate it from var. yunnanensis seem to be rather
similar with those of the latter instead. Here var. Pinus yunnanensis Franch. var. yunnanensis. Pinus
tenui­folia is considered synonymous with P. yun­ tabuliformis Carrière var. yunnanensis Dallim. &
nanensis var. yunnanensis. A. B. Jacks., Handb. Conif., ed. 3: 563. 1948; Pinus
insularis Endl. var. yunnanensis (Franch.) Silba,
Phytologia Mem. 7: 52. 1984; Pinus kesiya Royle ex
Distribution
Gordon subsp. yunnanensis (Franch.) Businský,
SW China: S Sichuan, Yunnan, E Guizhou, E Guang­ Acta Pruhoniciana 68: 24. 1999. Type: China: Yun-
xi. nan, Lijiang Xian, Dapingzi, [“bois des montagnes
TDWG codes: 36 CHC-GZ CHC-SC CHC-YN CHS- au dessus de Ta pin tze”], P. J. M. Delavay 569 (holo-
GX type P).
783
Pinus yunnanensis Franch. var. tenuifolia W. C. Cheng
Ecology
& Y. W. Law, Acta Phytotax. Sin. 13 (4): 85. 1975; Pi­
Pinus yunnanensis grows from 400 m to 3100 m nus insularis Endl. var. tenuifolia (W. C. Cheng &
a.s.l. in valleys and deep river gorges and on the Y. W. Law) Silba, Phytologia 68: 51. 1990.
high mountain slopes where it forms extensive for-
ests. It is most abundant on dry and sunny slopes
Description
and it can locally reach to the tree line, forming an
alpine shrub. It is adapted to survive on sites with Trees to 30 m tall. Leaves not or slightly pendulous,
infertile and shallow, rocky soils exposed to erosion. 10–20(–30) cm long, with 4–6 resin ducts. Seed cones
At high altitudes it can withstand severe frosts. It is opening at maturity, falling after seed ­dispersal.
most commonly gregarious in pure stands, but can
be found growing with other pines, e.g. P. armandii,
Distribution
P. tabuliformis and P. kesiya, or with Keteleeria evely­
niana, as well as with angiosperm trees and shrubs. SW China: S Sichuan, Yunnan, E Guizhou, E Guang­
xi.
TDWG codes: 36 CHC-GZ CHC-SC CHC-YN CHS-GX
Uses
Yunnan pine is of considerable economic im-
Conservation
portance in China, with natural stands covering
between 5.5 and 6 million ha and 275,000 ha in plan- IUCN: LC
tations. Its wood is used for fuel, as round wood for
transmission poles and stakes, as sawn timber for Pinus yunnanensis Franch. var. pygmaea (J. R. Xue)
construction, both exterior and interior, for fences J. R. Xue, Fl. Reipubl. Pop. Sin. 7: 258. 1978. Pinus
and gates, pallets, crates and boxes, cooperage and densata Mast. var. pygmaea J. R. Xue, Acta Phyto-
vats, furniture, veneers, all kinds of plywood and tax. Sin. 13 (4): 85. 1975; Pinus tabuliformis Carrière
boards, tool handles, pulp for paper, and to make var. pygmaea (J. R. Xue) Silba, Phytologia 68: 63.
chemical products like plastics and cellulose deriva- 1990. Type: China: Yunnan [locality not stated],
tives. Resin is tapped for naval stores, in particular Y. C. Hsüeh 156 (holotype PE).
turpentine, and the bark is used to extract tannins.
The leaves (needle litter) provide fodder for animals
Description
and are distilled for oils and medicinal products.
This species is much planted for wind breaks and A shrub-like high mountain form to 2 m tall,
erosion control. It is common in China as an orna- branching from base. Leaves 7–13 cm long, rigid, not
mental tree, but is not widely planted in other coun- drooping, with 2–3 resin ducts. Seed cones persist-
tries, where it is mostly seen in arboreta and other ent, more or less serotinous.
collections. It has a tendency to become invasive as
it is an aggressive pioneer species.
Distribution
2 varieties are recognized: China: SW Sichuan, Yunnan.
TDWG codes: 36 CHC-SC CHC-YN
 Ecology
This variety occurs in the mountains at altitudes be-
tween 2200 m and 3100 m a.s.l.

Conservation
IUCN: LC

784
 fi g u re 2 40 . Pinus figure 2 4 2. Pinus pinea in Algarve, Portugal
pinaster subsp. pinaster
pollen cones f i g u re 2 43. Pinus pinea seed cone

figure 2 45. Pinus

pungens seed cones

fi g u re 2 4 1 . Pinus pinaster subsp. pinaster

seed cones of two ages

f i g u re 2 4 4 . Pinus ponderosa
var. ponderosa pollen cones figure 2 46. Pinus quadrifolia in California, USA
 fig ure 2 4 8. Pinus rzedowskii
seed cones

fi g u re 2 4 7. Pinus rzedowskii tree figure 2 4 9. Pinus sylvestris

in Michoacán, Mexico var. sylvestris trees
figure 2 5 1. Pinus virginiana
seed cones

f i gu re 2 50 . Pinus taiwanensis var. taiwanensis in Hehuan Shan, Taiwan

figure 2 53. Platycladus

­orientalis foliage and seed
cones (photo D. Mabberley)
figure 2 54. Platycladus
­orientalis seed cones

figure 2 52. Pinus

­wallichiana var. wallichiana
seed cones
f i gu re 2 5 5 . Podocarpus acutifolius foliage
and pollen cones

figure 2 5 7. Podocarpus brevifolius on

the Mesilau River, Mt. Kinabalu, Borneo

figure 2 56. Podocarpus brassii var.

brassii seed cone (photo T. Waters)

figure 2 58. Podocarpus

brevifolius leaves

figure 2 59. Podocarpus chingianus

­foliage and seed cone

figure 2 61. Podo­

carpus cunninghamii
foliage and seed cones

figure 2 62. Podo­

carpus dispermus
leaves

figure 2 60. Podo­

carpus costalis foliage
 fig ure 2 63. Podocarpus
elatus foliage

fig ure 2 64. Podocarpus elatus

seed cone

figure 2 65. Podocarpus grayae

small sapling at Cape Tribulation,
Queensland

fig ure 2 6 9. Podocarpus

­laubenfelsii seedling
on Mt. Kinabalu

figure 2 68. Podocarpus

fi g u re 2 66. Podocarpus grayae tree at henkelii leaves
Lake Eacham, Queensland
figure 27 0. Podocarpus
l­atifolius in the Drakensberg, S.A.
(photo J. Grimshaw)

figure 2 67. Podocarpus

grayae trunk, Herberton Range,
Queensland
 figure 27 7. Podocarpus nakaii foliage
and seed cones
f i gu re 2 7 2 . Podocarpus macrophyllus
var. macrophyllus foliage figure 27 1. Podocarpus lawrencei
seed cones
figure 27 6. Podocarpus
nakaii flushing leaves

figure 27 3. Podocarpus
macrophyllus var. macrophyllus
foliage and pollen cones
figure 279. Podocarpus
neriifolius var. neriifolius seed
cones (photo T. Utteridge)

figure 274. Podocarpus

matudae leaves and young
seed cone
figure 27 8. Podocarpus
neriifolius var. neriifolius in
P.N.G. (photo T. Utteridge)

figure 27 5. Podocarpus
­milanjianus on Mt. Elgon,
Uganda (photo D. L. Roberts)
 figure 2 83. Podocarpus novae-caledoniae
foliage and young seed cones
figure 2 80. Podocarpus nivalis in North Island,
New Zealand
figure 2 81. Podo­
carpus nivalis foliage
and seed cones

figure 2 84. Podo­

carpus nubigenus
leaves (upperside)

figure 2 86. Podocarpus

­polystachyus pollen cones

f i g ure 2 85. Podocarpus

­nubigenus leaves (underside)

fig ure 2 82. Podocarpus

­novae-caledoniae in
New Caledonia

figure 2 87. Podocarpus rumphii seed cones

fi g u re 2 8 8 . Podocarpus fig ure 2 89. Podocarpus spinulosus on North Stradbroke Island, Queensland
salignus foliage and young
seed cones

fi g u re 29 5 . Podocarpus totara foliage

and pollen cones

figure 290. Podocarpus spinulosus figure 2 9 1. Podocarpus

unripe seed cone (photo G. Garruthers) spinulosus ripe seed cones
(­photo G. Garruthers)
fig ure 293. Podocarpus totara tree in North Island, New Zealand
figure 292. Podocarpus sprucei foliage buds figure 2 94 . Podo-
(photo P. Cribb 1425) carpus totara bark
 f i gu re 29 6 . Prumnopitys andina foliage and figure 297. Prumnopitys andina seed cones
pollen cones

figure 299. ­Prumnopitys

­ladei trunk on Mt. Lewis,
Queensland

fi g u re 29 8 . Prumnopitys ferruginoides foliage

on Mt. Mou, New Caledonia

f i gu re 3 0 0 . Prumnopitys ladei foliage

figure 301. Pseudolarix amabilis
pollen cones

figure 303. Pseudotaxus chienii foliage and

fi g u re 3 02 . Pseudolarix amabilis seed cones buds of seed cones
Platycladus Spach, Hist. Nat. Vég. Phan. 11: 333. 1841. Type: Platycladus orientalis
(L.) Franco [Platycladus stricta Spach (nom. illeg.) (Thuja orientalis L.)] (Cupressa­
ceae).

Biota (D. Don) Endl., Syn. Conif.: 46. 1847, non Cas- dères & Gaussen) A. V. Bobrov, 14. Symp. Biodiv.
sini (1825). Type: Biota orientalis (L.) Endl. [Platy­ Evol. Biol. Jena 1999: 18. 1999.
cladus orientalis (L.) Franco (Thuja orientalis L.)]
Etymology
Greek: platys = broad, flat; klados = branch; referring
to the flattened foliage branches. The species epithet means ‘from the east’, wherewith 793
Linnaeus distinguished the species from his other
species in Thuja.
Description
See the species description.
Vernacular names
Oriental Arbor-vitae, Chinese Arbor-vitae; ce bai
Distribution
(Chinese)
As for the species.
Description
Taxonomic notes
Trees to 20–25 m tall, evergreen, monoecious; trunk
Detailed observations of the early stages of seed cone monopodial or multistemmed from low above
development in both Microbiota and Platycladus ground, to 2–2.5(–4) m d.b.h., with stem(s) erect or
(Jagel & Stützel, 2001) demonstrated that these taxa ascending. Bark on large stems becoming fibrous,
are probably closely related. Both cones are quite exfoliating in thin, long sheets, dull red-brown.
variable (variation is greater in Platycladus) and are Branches spreading, ascending or erect, contorted in
primarily distinct in size and in the number of bract- old trees, higher order branches mostly spreading,
scale complexes and ovules. Extremely reduced forming a pyramidal, rounded or irregular crown.
cones of Platycladus resemble strongly developed Foliage branches in flattened, more or less vertically
cones of Microbiota; the latter seems merely a cone of disposed sprays, in older trees more horizontal or
Platycladus strongly reduced in size and number of irregularly disposed, persistent. Leaves decussate,
parts, including seeds. Phylogenetic analyses of the decurrent, scale-like, on lateral branchlets dimor-
genera in Cupressaceae (Brunsfeld et al., 1994; Gadek phic, facials more or less rhombic to obtrullate,
et al., 2000) based primarily on molecular data, con- with appressed, obtuse apex; laterals bilaterally flat-
firmed the close relationship of these two taxa and tened, distal part spreading or reflexed, appressed or
their much greater distance from Thuja. A concise free, with incurved, obtuse apex; leaves on ultimate
review of Platycladus, with botanical illustrations, branchlets 1.5–2 × 1–1.5 mm, with facials slightly
was published by Morgan (1999) in Curtis’s Botanical smaller than laterals; amphistomatic, stomata in
Magazine, hopefully with the effect that this genus irregular lines; gland more or less linear, inactive
will be recognized by all as truly distinct from Thuja. or sometimes active; leaf colour green or greyish
green. Pollen cones terminal on ultimate branchlets,
solitary, subglobose, 2–3 × 2 mm; microsporophylls
Platycladus orientalis (L.) Franco, Portugaliae Acta (6–)8–10(–12), decussate, peltate, bearing 3–6 abax-
Biol., sér. B, Sist. Vol. “Julio Henriques”: 33. 1949. ial pollen sacs. Seed cones terminal, maturing in
Thuja orientalis L., Sp. Pl. 2: 1002. 1753. Type: China: one year to erect, ampulliform, glaucous or purplish
[locality unknown], leg. ign. LINN 1136.2 (lectotype cones 15–20(–25) × 10–18 mm which open widely
LINN). Fig. 253, 254 and turn brown when the seeds have ripened. Bract-
scale complexes 6–8, decussate, with mostly sterile
Thuja chengii Bordères & Gaussen, Bull. Soc. Hist. upper pair, oblong to obtrullate, variable in size up
Nat. Toulouse 73: 284. 1939; Platycladus chengii (Bor- to 15 × 8 mm, with a prominently recurved ­subapical
 spine 3–8 mm long, spreading wide, more or less climax. The climate in NE China is above all char-
woody, thick, exposing often persistent seeds. Seeds acterized by very cold winters. As a pioneer of rela-
6–10(–12?), ovoid to narrowly ovoid, 5–7 × 3–4 tively dry, open vegetation on often unstable slopes,
mm, greyish brown with a large, light brown hilum; P. orientalis has found abundant opportunity over
wings absent or rudimentary as two narrow strips much of China, and even beyond (e.g. NE Iran), to
near seed apex. establish itself and spread after introduction. It is
much used in afforestation in NE and Central China
and commonly planted in Central Asia. Buddhism
Distribution
has been instrumental in its spread especially in the
China: S Gansu, Hebei, Henan, Shaanxi, Shanxi; SW of China, where it can be abundant on steep
794 Korea; Russian Far East. [This is possibly the most slopes of river valleys, but is usually never very far
widely introduced cupressaceous conifer in Asia. In from settlements, monasteries or temples.
many areas inside and outside China it has ‘escaped’
from cultivation and established spontaneous popu-
Conservation
lations. Only its natural (indigenous) distribution is
given here.] IUCN: NT
TDWG codes: 31 AMU KHA 36 CHN-GS CHN-HB
CHN-SA CHN-SX CHS-HE 38 KOR-NK KOR-SK
Uses
The wood of Oriental Arbor-vitae is used for build-
Ecology
ing and construction of houses and temples where
Platycladus orientalis is most probably a species of there are still trees of good size left. The foliage is in
the transitional open woodland zone between the some parts of China much used for incense burning,
steppes of Inner Mongolia and the deciduous oak, to which purpose the species has been introduced
oak-birch, and oak-pine forests of NE China. Even widely outside its natural range. It is also one of the
within its natural range it is now almost invariably most commonly planted amenity and ornamental
found in secondary vegetation or, nearest to its orig- conifers, a tradition that goes back many centuries.
inal habitat, in more or less degraded woodland and It is therefore a common tree in parks of towns and
forest. As a pioneer species which is relatively long- cities in much of temperate Asia; it appears to be
lived, it can dominate certain slopes for a long time tolerant of drought as well as the air pollution that
if further disturbances remain absent. Elsewhere occurs within the urban environment. In some parts
it grows together with Pinus tabuliformis, less fre- of China (e.g. Yunnan) and in other countries (e.g.
quently with P. armandii and Juniperus rigida; Betu­ Iran) it has probably been naturalized. In NE China
la chinensis and Populus tremula are followed in the it is being used in afforestation schemes on defor-
succession by Quercus spp. (some of which may be ested hills and mountains. A substantial number of
evergreen) and Castanea. The most arid and steepest cultivars has been raised in Europe since its intro-
slopes may only be covered with Pinus tabuliformis duction to France around 1700, many of which are
and Platycladus, accompanied by Juniperus rigida now obsolete, while new cultivars continue to be se-
and other shrubs, and this vegetation may not rep- lected and described.
resent a seral type, but an edaphically determined
Podocarpus L’Hér. ex Pers., Syn. Pl. 2 (2): 580. 1807 (nom. cons.). Type: Podocarpus
elongatus (Aiton) L’Herit. ex Pers. [Taxus elongata Aiton] (Podocarpaceae).

Margbensonia A. V. Bobrov & Melikyan, Byull. and southern Japan; from Indochina throughout
Moskovsk. Obshch. Isp. Prir., Otd. Biol. 103 (1): 59. Malesia (excluding Lesser Sunda Islands) to New
1998. Type: Margbensonia macrophylla (Thunb.) Britain. Australia: SW corner of Western Australia,
A. V. Bobrov & Melikyan [Podocarpus macrophyllus Northern Territory (Arnhem Land), Queensland,
(Thunb.) Sweet (Taxus macrophylla Thunb.)] New South Wales, Victoria, Tasmania. SW Pacific:
Solomon Islands, New Caledonia, Vanuatu, Fiji,
Greek: podo = foot, carpus = fruit; alluding to the Tonga, New Zealand. North America: S Mexico. 795
receptacle and seed respectively of the female repro- Caribbean Islands (excluding Bahamas). Central
ductive organ (cone + seed). America (all countries). South America: Colombia,
Venezuela, Guyana, Ecuador, Brazil, Bolivia, N Ar-
gentina, S Chile.
Description
(Decumbent) shrubs or more commonly trees, dio-
Taxonomic notes
ecious or rarely monoecious, much branched. Bark
scaly or fibrous, peeling in vertical strips. Resin ca- The taxonomy of the genus Podocarpus, after Pi­
nals mostly in the leaves, none in the wood. Primary nus the most speciose (but not the most diverse)
branching in pseudo-whorls, plagiotropic (Massart’s genus in the conifers, is perhaps the least well set-
model). Terminal buds distinctive, with imbricate tled among the major coniferous genera. Whereas
and/or spreading primary and secondary scales. at the family level a limited number of phylogenetic
Leaves spirally inserted or subopposite on lateral fo- studies have now been published, which appear
liage branchlets, relatively broad, bifacially flattened, to mostly corroborate the narrower circumscrip-
usually linear-lanceolate or linear-elliptic, sessile to tion of the genus established in recent decades on
short petiolate, coriaceous, with a single raised, flat morphological grounds, no such studies have been
or sunken midrib and stomata in two broad bands attempted that more or less sampled the genus Podo­
on abaxial side, rarely on both sides and then only carpus comprehensively. There are two revisions of
few stomata on adaxial side. Pollen cones axillary, the genus based on anatomy and/or morphology
solitary or clustered, sessile or on short peduncles, that need to be briefly assessed here, in order to es-
each cone subtended by bud scales, flexible and tablish whether they are useful for this Handbook.
slender, catkin-like. Microsporophylls spirally at- The first is by J. T. Buchholz and N. Gray and was
tached to a slender rachis, triangular, with two basal published in a series of papers (variously authored
pollen sacs containing bisaccate pollen. Seed cones by both or by Gray alone later) in the Journal of
axillary, pedunculate, solitary or rarely more togeth- the Arnold Arboretum in 1948–1962. The circum-
er, consisting of 2–5 adnate bracts (upper 1–2 fertile); scription of Podocarpus still included several other
the sterile bracts fusing and greatly swelling to form genera (as sections) and only their section “Eupodo­
a smooth, succulent, coloured receptacle. Seeds 1, carpus” (an illegitimate name under the Botanical
sometimes 2, obliquely attached to the apical part of Code) was equivalent to the genus Podocarpus as
the receptacle, completely exposed, morphologically presently understood. This section was divided into
inverted, drupe-like, completely covered by a fleshy, six subsections, informally named A-F. The empha-
green or sometimes reddish to bluish epimatium. sis was on a detailed study of leaf anatomy, but the
subsections were mostly defined by geography and
97 species. a few selected characters, indeed mostly anatomi-
cal. De Laubenfels (1985) followed an earlier sug-
gestion by Pilger (1926) that the presence or absence
Distribution
of ‘foliola’ (lanceolate bracts) below the receptacle
Africa: S Nigeria, Cameroon, Angola; in E Africa could be used to subdivide the genus Podocarpus
from extreme S Sudan to South Africa down to the (now in the narrower sense still in use) into two
Cape; Madagascar. Asia: from E Nepal to Taiwan subgenera. Further “special characteristics” then
 helped De Laubenfels to further subdivide the genus 4a. Seeds including the epimatium ca. 8 × 5 mm.
into 18 sections “of from 1–10 species each” and the Scales of terminal buds obtuse or acute. Leaves
two subgenera and 18 sections were then formally 7–20(–25) mm long P. perrieri
named and described. In a phylogenetic context, it 4b. Seeds including the epimatium ca. 14 × 8 mm.
is unlikely that such a scheme, based as it is on a few Scales of terminal buds apiculate. Leaves (10–)
selected characters, would be supported by a cladis- 15–30(–36) mm long P. rostratus
tic analysis of these and other anatomical or mor- 5a. Leaves 2–5 mm wide, up to 7.5 cm long (mature
phological characters or of molecular data. Simply trees!). Bud scales elongated P. capuronii
put, a different classification would result from the 5b. Leaves 3–16 mm wide, up to 18 cm long (but
choice of different characters, and without proper highly variable!). Bud scales rounded to ovate
796 testing we do not know how ‘artificial’ or ‘natural’ P. madagascariensis
such a scheme might be. It must therefore be con- 6a. Midrib on adaxial (upper) side of leaves dis-
cluded that neither of these two classifications has tinct and continuously raised. Pollen cones al-
been tested, either by repeated (phenetic or phyletic) ways solitary P. milanjianus
attempts using additional characters (as has been 6b. Midrib on adaxial (upper) side of leaves indis-
the case with Abies and Pinus) or by a comprehen- tinct and/or fading towards the apex. Pollen
sive cladistic analysis of similar data or molecular cones 2–5 together, rarely solitary 7
data (as with Abies and especially Pinus). No clas- 7a. Shrubs or small trees to 6 m tall (occasionally
sification of Podocarpus is therefore presented in to 20 m). Leaves (3–)4–5(–7) mm wide
this Handbook. The large number of species and the P. elongatus
paucity of useful morphological characters make it 7b. Trees to 30–35 m tall (occasionally stunted on
impractical to construct a single key to all species. exposed mountain tops). Leaves (5–)6–12(–18)
I therefore present keys to species of Podocarpus mm wide 8
grouped on a geographical basis. An additional key 8a. Apex of leaves acute or acuminate. Pollen cones
is presented to help identify species that are more or (1–)2–5 together. Receptacle below the seed ru-
less commonly found in cultivation. dimentary; seeds including the epimatium 17–
22 mm long P. henkelii
8b. Apex of leaves obtuse or mucronate. Pollen
Key to the species of Podocarpus in Africa
cones 1–2 together. Receptacle below the seed
and Madagascar
8–14 × 8–12 mm, succulent; seeds including the
When using this key in the field only leaves on epimatium 7–11 mm long P. latifolius
branches of mature shrubs or trees, preferably from
sun-exposed foliage, should be taken into account.
Key to the species of Podocarpus in China
Male (pollen) and female cones and seeds (they oc-
(including Taiwan), NE India, Indochina,
cur on dioecious shrubs and trees) are described as
and Japan
mature.
When using this key in the field only leaves on
1a. Scales of terminal buds all imbricate or in- branches of mature trees, preferably from sun-­
curved 2 exposed foliage, should be taken into account. Male
1b. Scales of terminal buds with at least some outer (pollen) and female cones and seeds (they occur on
scales spreading or recurved 6 dioecious trees) are described as mature.
2a. Leaves 7–15 mm long, 2–4 mm wide, obovate to
oblanceolate (widest above the middle) 1a. Buds globose to short conical, with imbricate,
P. humbertii triangular scales 2
2b. Leaves usually longer than 15 mm (if smaller, 1b. Buds variously shaped but not globose, with
only 1–2 mm wide), linear or widest below the free, erect to spreading or recurved (outer)
middle 3 cales 3
3a. Leaves 7–36 mm long, 1–2 mm wide, narrowly 2a. Adult leaves (3–)5–10(–12) cm long, lanceolate
lanceolate to linear 4 to linear-lanceolate; midrib on adaxial (upper)
3b. Leaves (1.5–)2–18 cm long, 2–16 mm wide, el- side of leaves acutely raised P. nakaii
liptic-oblong to linear 5
2b. Adult leaves 9–14(–22) cm long, linear; midrib as far as New Guinea and the main chain of the Solo-
on adaxial side of leaves obtusely raised mon Islands. Species which occur only in Borneo or
P. rumphii New Guinea including the Bismarck ­Archipelago
3a. Adult leaves 1.2–5(–7) cm long. Bud scales erect and/or the Solomon Islands are dealt with in two
or slightly preading 4 keys specific to these areas; species which are com-
3b. Adult leaves (2.5–)4–12(–15) cm long. Bud mon to two or more of these areas appear conse-
scales spreading or recurved, sometimes more quently in more than one key. When using this key
or less erect 6 in the field only leaves on branches of mature trees,
4a. Adult leaves 2.5–4.5 mm wide, not longer than preferably from sun-exposed foliage, should be tak-
3.5 cm P. chingianus en into account. Male (pollen) and female cones and
4b. Adult leaves (4–)5–8(–10) mm wide, often seeds (they occur on dioecious trees) are described 797
longer than 3.5 cm (maximum length 5–7 cm) as mature.
5
5a. Pollen cones solitary, sessile, 8–25 × 6–8 mm. 1a. Buds short conical or subglobose; bud scales
Small trees 1.5–10 m tall P. costalis imbricate 2
5b. Pollen cones solitary, pedunculate, 15–30 × 3 1b. Buds elongated, acute; outer bud scales free,
mm. Shrubs or trees to 25 m tall P. pilgeri erect or spreading 4
6a. Bud scales caudate, recurved. Midrib on adax- 2a. Bud scales rounded at apex. Midrib of leaves on
ial (upper) side of leaves obtusely raised adaxial (upper) side acutely raised, narrow, less
P. macrophyllus than 1 mm wide P. teysmannii
6b. Bud scales triangular-lanceolate to lanceolate, 2b. Bud scales triangular, with an acute apex. Mid-
erect, spreading or sometimes recurved. Mid- rib of leaves on adaxial side obtusely raised,
rib on adaxial side of leaves acutely raised 7 1 mm wide 3
7a. Buds 1.5–3 mm long. Adult leaves from nearly 3a. Buds up to 4 × 4 mm, short conical. Seeds in-
oval to linear, with an acute or obtuse apex cluding the epimatium ca. 10 mm long
P. polystachyus P. laubenfelsii
7b. Buds (2–)3–5(–8) mm long. Adult leaves lan- 3b. Buds (2.5–)4–5(–8) mm long, globose to short
ceolate to linear-lanceolate, with an acute or conical. Seeds including the epimatium 12–15
acuminate apex 8 mm long P. rumphii
8a. Pollen cones 15–25(–30) × 2–2.5 mm. Recepta- 4a. Adult leaves 0.8–3(–3.5) cm long, 3–6 mm wide.
cles when ripe 9–16 mm long; seeds including Shrubs or small trees 5
the epimatium ca. 9 × 7 mm P. fasciculus 4b. Adult leaves mostly longer. (Small) trees 6
8b. Pollen cones 20–40(–60?) × 2–3.5 mm. Recep- 5a. Adult leaves elliptic to obovate; apex obtuse to
tacles when ripe 7–12 mm long; seeds including abruptly rounded P. glaucus
the epimatium (8–)10–15 × 8–10 mm 9 5b. Adult leaves elliptic to oblong; apex acute to ob-
9a. Receptacles subtended by two 2–6 mm long tuse P. lophatus
bracts (‘foliola’); epimatium around the seeds 6a. Adult leaves (4–)5–8(–10) mm wide 7
without a crest. New leaves flushing red 6b. Adult leaves (6–)8–18(–20) mm wide 10
P. neriifolius 7a. Buds 6–14 mm long; outer bud scales erect with
9b. Receptacles subtended by two 1–2 mm long recurved tips. Adult leaves 6–11 cm long
bracts; epimatium around the seeds with a P. atjehensis
crest. New leaves flushing yellowish green 7b. Buds 2–5 mm long; outer bud scales erect or
P. subtropicalis spreading, with straight tips. Adult leaves (2–)
3–6(–7) cm long 8
8a. Pollen cones solitary, sessile, 6–8 mm diam
Key to the species of Podocarpus in Malesia,
P. costalis
excluding Borneo and New Guinea, and
8b. Pollen cones solitary or more often with 2–3
including the Andaman Islands (India)
together, pedunculate when solitary, otherwise
Malesia is the phytogeographic region as defined in sessile, 2.5–3.5 mm diam 9
Flora Malesiana, and includes Peninsular Malaysia 9a. Apices of adult leaves acute. Pollen cones
and Singapore and all the islands of the archipelago solitary or often with 2–3 together, sessile.
 ­ eceptacles 6–8 mm long when ripe; seeds in-
R including the epimatium 6 × 5 mm, purple
cluding the epimatium 8–9 × 5–6 mm, red or P. borneensis
purple P. rubens 18b. Adult leaves (4–)8–12(–15) cm long, with an
9b. Apices of adult leaves of various shapes, but acute apex. Receptacles 8–14 mm long; seeds
often obtuse to rounded. Pollen cones solitary, including the epimatium (8–)10–15 × 7–8 mm
pedunculate. Receptacles 10–22 mm long when 19
ripe; seeds including the epimatium 10–12 × 19a. Buds 2–5 mm long, sometimes with a few long-
8–9 mm P. pilgeri er outer scales to 8 mm. Receptacles when ripe
10a. Adult leaves (1.5–)2.5–5 cm long, elliptic to obo- 8–10 mm long P. neriifolius
vate, with an obtuse or rounded apex P. ramosii 19b. Buds 5–12 mm long, with all or most of the
798 10b. Adult leaves (2.5–)4–15(–18.5) cm long, nearly outer scales the same length. Receptacles when
oval to linear, rarely elliptic, with an acute, ob- ripe 10–14 mm long P. bracteatus
tuse or rounded apex 11
11a. Leaves linear, deflected at the petiolate base Key to the species of Podocarpus in Borneo
P. deflexus
When using this key in the field only leaves on
11b. Leaves nearly oval to linear-lanceolate or linear,
branches of mature trees, preferably from sun-­
not deflected 12
exposed foliage, should be taken into account. Male
12a. Adult leaves (8–)12–20 mm wide (5–10 cm
(pollen) and female cones and seeds (they occur on
long), with obtuse or rounded apex
dioecious trees) are described as mature.
P. spathoides
12b. Adult leaves 6–15(–18) mm wide, with acute or 1a. Bud scales imbricate 2
obtuse apex (if wider than 14 mm, apex acute) 13 1b. Bud scales free, erect or spreading 5
13a. Midrib on adaxial (upper) side of leaves indis- 2a. Buds 2–5 mm long, scales rounded 3
tinct, narrow; adult leaves very gradually taper- 2b. Buds 4–8 mm long, scales triangular 4
ing to a narrowly acute apex. Receptacle pink 3a. Buds 2–3 mm long. Leaves (2.5–)4–8(–9) cm
when ripe P. ridleyi long, (7–)9–15 mm wide P. globulus
13b. Midrib on adaxial side of leaves distinct, acutely 3b. Buds 3–5 mm long. Leaves 7–14 cm long, (10–)
or obtusely raised; adult leaves acute or obtuse, 12–17(–20) mm wide P. teysmannii
sometimes with a rounded apex. Receptacle red 4a. Bud scales with recurved apex. Pollen cones
when ripe 14 3–5 together, short pedunculate P. laubenfelsii
14a. Pollen cones solitary, 6.5–8 mm wide. Small 4b. Bud scales entirely appressed. Pollen cones (2–)
trees as far as known P. palawanensis 3–5 together, sessile P. rumphii
14b. Pollen cones solitary or more often with 2–3 5a. Leaves 1–7 cm long, rarely to 9 cm long, 3–13
(–5) together, 2–4 mm wide. Potentially large mm wide; shorter leaves usually elliptical,
trees 15 sometimes obovate; longer leaves oblong or
15a. Midrib on adaxial (upper) side of leaves ob- linear-lanceolate 6
tusely raised, fading towards the apex 16 5b. Leaves (2.5–)5–15(–18) cm long, (5–)6–20 mm
15b. Midrib on adaxial side of leaves acutely raised, wide, usually linear-lanceolate, sometimes lin-
always continuous to the apex 17 ear-oblong; shorter leaves sometimes elliptical
16a. Buds 2–4 mm long. Pollen cones to 25 mm long. or nearly oval 11
Receptacles 10–15 mm long; seeds including the 6a. Buds 2–5 mm long 7
epimatium 14–17 × 10–13 mm P. macrocarpus 6b. Buds 4–10 mm long 9
16b. Buds 3–9 mm long. Pollen cones 50–60(–80?) 7a. Buds 2–3 mm long. Leaves elliptical to obovate,
mm long. Receptacles 8 mm long; seeds includ- 7–13 mm wide; apex obtuse or rounded
ing the epimatium 8–11(–13?) × 7–8 mm P. levis P. ramosii
17a. Buds 1.5–3 mm long. Pollen cones in clusters of 7b. Buds 2–5 mm long. Leaves linear-elliptic to
2–5, 15–30 mm long P. polystachyus linear-lanceolate, (4–)5–8(–10) mm wide; apex
17b. Buds 2–12 mm long. Pollen cones solitary or acute, obtuse, or rounded 8
with 2–3 together, 25–50(–60) mm long 18 8a. Leaves flushing green. Pollen cones always soli-
18a. Adult leaves 3–7(–9) cm long, with an obtuse tary, pedunculate. Receptacle when ripe 10–22
or rounded apex. Receptacles 6 mm long; seeds mm long P. pilgeri
8b. Leaves flushing red. Pollen cones solitary or of- 15b. Midrib on adaxial (upper) side of leaves usually
ten 2–3 together. Receptacle when ripe 6–8 mm acutely raised, continuous to the acute apex.
long P. rubens Seeds including the epimatium (8–)10–15 mm
9a. Bud scales acuminate, spreading. Pollen cones long P. neriifolius
solitary, rarely in pairs, ca. 15 × 3–4 mm
P. gibbsiae
Key to the species of Podocarpus in Papuasia
9b. Bud scales triangular or lanceolate-linear, erect
or spreading. Pollen cones solitary or often 2–3 Papuasia is the phytogeographic region that in-
together, 20–50 × 2.5–5 mm 10 cludes New Guinea, the Bismarck Archipelago and
10a. Bud scales 4–6 mm long, triangular, spreading. the Solomon Islands (as on Plate 12 of the Times
Pollen cones solitary or in pairs, ca. 20 × 4–5 Atlas, 12th edition, but excluding the Santa Cruz Is- 799
mm P. brevifolius lands). When using this key in the field only leaves
10b. Bud scales 4–10 mm long, lanceolate-linear, on branches of mature trees, preferably from sun-
erect or slightly spreading. Pollen cones solitary exposed foliage, should be taken into account. Male
or with 2–3 together, 35–50 × 2.5–3.5 mm (pollen) and female cones and seeds (they occur on
P. borneensis dioecious trees) are described as mature.
11a. Bud scales triangular, spreading. Leaves linear-
oblong, shortest ones elliptical; apex obtuse or 1a. Usually shrubs, only rarely (small) trees. Adult
rounded P. spathoides leaves not longer than 3 cm 2
11b. Bud scales narrowly triangular to lanceolate- 1b. Trees (can be stunted at high altitudes). Adult
linear or long acuminate, erect or spreading. leaves usually longer than 3 cm 3
Leaves linear-lanceolate, shortest ones lanceo- 2a. Buds 3–5 mm long. Pollen cones ca. 40 × 3–8
late or nearly oval 12 mm. Receptacles when ripe 12–15 mm long,
12a. Bud scales long acuminate, erect. Pollen cones thick, purple or purple-black; seeds including
solitary, rarely in pairs. Receptacle with a single the epimatium 10–13 × 8–9 mm, without an
basal bract 3–5 mm long P. confertus apical crest P. brassii
12b. Bud scales triangular to lanceolate-linear, erect 2b. Buds 2–3 mm long. Pollen cones 10–25 × 6–7
or spreading. Pollen cones solitary or more of- mm. Receptacles when ripe 6–8 mm long,
ten with 2–5 together. Receptacle with 2 basal purple; seeds including the epimatium 6–7 × 5
bracts 1.5–6 mm long 13 mm, with a small apical crest P. glaucus
13a. Buds 1.5–3 mm long. Leaves nearly oval to lin- 3a. Adult leaves (1.5–)2.5–6 cm long 4
ear-lanceolate. Pollen cones 2–5 together, 15–30 3b. Adult leaves mostly longer than 5 cm, only a
× 3 mm. Seeds including the epimatium 7–9 × few sometimes shorter 5
5–7 mm P. polystachyus 4a. Pollen cones always solitary and pedunculate.
13b. Buds (2–)3–15 mm long. Leaves linear-lanceo- Receptacles when ripe 10–22 mm long; seeds
late or lanceolate. Pollen cones solitary or 2–3 including the epimatium 10–12 × 8–9 mm
together, (25–)30–75(–80?) × 2–4 mm. Seeds in- P. pilgeri
cluding the epimatium (6–) 8–15 × 6–8 mm 14 4b. Pollen cones solitary or often 2–3 together, ses-
14a. Buds 6–15 mm long, scales long linear-lanceo- sile. Receptacles when ripe 6–8 mm long; seeds
late, erect or spreading. Leaf apex acuminate or including the epimatium 8–9 × 5–6 mm
acute. Epimatium with a distal crest P. rubens
P. micropedunculatus 5a. Buds globose to conical, as wide as long, with
14b. Buds (2–)3–9 mm long, scales lanceolate or imbricate scales. Leaves all linear P. rumphii
rarely triangular, spreading. Leaf apex acute 5b. Buds variously shaped but not globose, includ-
or obtuse. Epimatium smooth without a distal ing the free, erect or spreading outer scales
crest 15 longer than wide. Leaves variously shaped, not
15a. Midrib on adaxial (upper) side of leaves ob- all linear 6
tusely raised, often fading towards the acute 6a. Leaves (strongly) deflected at the petiolate
or obtuse apex. Seeds including the epimatium base P. atjehensis
8–11 mm long P. levis 6b. Leaves spreading or sometimes drooping, not
deflected 7
 7a. Apex of adult leaves obtuse or rounded, leaves 15a. Buds 2–5 mm long, sometimes with a few long-
5–10 cm long, (8–)12–20 mm wide. Receptacles er outer scales to 8 mm. New leaves flushing
when ripe 5 mm long; seeds including the epi- red P. neriifolius
matium 7 × 5 mm, purple P. spathoides 15b. Buds 2–3 mm long, without longer outer scales.
7b. Apex of adult leaves acute, acuminate or ob- New leaves flushing pale whitish green
tuse, never rounded, if adult leaves 20 mm P. insularis
wide (or wider), longer than 10 cm. Receptacles
when ripe longer than 8 mm; seeds including
Key to the species of Podocarpus in Australia
the epimatium larger than 7 × 5 mm, variously
(purplish) green 8 When using this key in the field only leaves on
800 8a. Midrib on adaxial (upper) side of leaves ob- branches of mature trees, preferably from sun-­
tusely raised, often fading towards apex 9 exposed foliage, should be taken into account. Male
8b. Midrib on adaxial (upper) side of leaves acutely (pollen) and female cones and seeds (they occur on
raised, continuous to apex 10 dioecious trees) are described as mature.
9a. Pollen cones always solitary, 30–40 × 6–8 mm.
Buds 3–5 mm long, with triangular to lanceo- 1a. Shrubs. Adult leaves 2–5 mm wide 2
late, slightly spreading scales. Seeds including 1b. Trees. Adult leaves (5–)7–30 mm wide 4
the epimatium 5–6 mm wide P. archboldii 2a. Adult leaves 0.4–1.6 cm long; leaf apices obtuse.
9b. Pollen cones often 2–3 together or solitary, 50 Bud scales imbricate, broadly triangular
(–80?) × 2–3.5 mm. Buds 3–9 mm long, with P. lawrencei
lanceolate, spreading outer scales. Seeds in- 2b. Adult leaves 2.5–9(–13) cm long; leaf apices
cluding the epimatium 7–8 mm wide P. levis acuminate or pungent. Bud scales free, (slight-
10a. Buds 6–14 mm long; bud scales with a long, ly) spreading, narrowly triangular 3
acuminate or caudate apex 11 3a. Apices of leaves apiculate, pungent; midrib on
10b. Buds 1.5–7(–8) mm long; bud scales triangular adaxial (upper) side of leaves acutely raised.
to lanceolate 13 Pollen cones 4–8 mm long. Receptacles when
11a. Adult leaves (11–)15–25 mm wide. Receptacles ripe 6–10 × 6–7 mm; seeds including the epi-
when ripe 10–16 mm long P. ledermannii matium 8–10 × 5–7 mm P. spinulosus
11b. Adult leaves (5–)7–12 mm wide. Receptacles 3b. Apices of leaves acuminate; midrib on adaxial
when ripe 8–11 mm long 12 side of leaves obtusely raised. Pollen cones (5–)
12a. All bud scales erect. Adult leaves (4–)6–15 cm 10–20(–22) mm long. Receptacles when ripe
long, leaf apices acute P. pseudobracteatus 20–25 × 10–13 mm; seeds including the epi­
12b. Outer bud scales spreading. Adult leaves 10–25 matium 12–20 × 8–12 mm P. drouynianus
cm long, leaf apices acute or obtuse 4a. Bud scales free, spreading. Adult leaves (13–)
P. salomoniensis 18–30 mm wide, oval to oblong P. dispermus
13a. Buds on leading shoots (3–)4–8 mm wide and 4b. Bud scales imbricate, appressed. Adult leaves
4–7 mm long, onion-shaped or truncate, with (5–)7–18(–20) mm wide, linear to linear-­
strongly recurved outer scales. Pollen cones lanceolate 5
6–7 mm wide P. crassigemma 5a. Adult leaves (6–)10–25(–30) cm long, linear
13b. Buds on leading shoots longer than wide, more P. grayae
or less conical, with erect or spreading outer 5b. Adult leaves (3–)5–11(–15) cm long, linear-­
scales. Pollen cones 2–3.5 mm wide 14 lanceolate 6
14a. Adult leaves 6–11 mm wide. Pollen cones al- 6a. Buds 1.5–2.5 mm long. Midrib on adaxial (up-
ways clustered with 2–5 together, 15–30 mm per) side of leaves obtusely raised but promi-
long. Seeds including the epimatium 7–9 mm nent. Pollen cones 10–20 × 2–3 mm. Receptacles
long P. polystachyus when ripe 15–25(–30) × 15–20 mm, dark purple
14b. Adult leaves 8–15(–18) mm wide. Pollen cones or blue-black P. elatus
solitary or often 2–3 together, 25–40(–50) mm 6b. Buds 3.5–8 mm long. Midrib on adaxial side of
long. Seeds including the epimatium (8–)10–15 leaves obscure. Pollen cones 30–45 × 4–6 mm.
mm long 15 Receptacles when ripe 5–7 × 5 mm, red
P. smithii
 8a. Pollen cones always solitary, 3.5–5 mm wide.
Key to the species of Podocarpus in the
Receptacles when ripe 10–13 × 8–9 mm. Adult
SW Pacific
leaves 6–8 mm wide P. polyspermus
The SW Pacific for the purposes of this key includes 8b. Pollen cones with 2–3 together or sometimes
New Caledonia, Vanuatu, Fiji, and Tonga. When us- solitary, 2.5–3.5 mm wide. Receptacles when
ing this key in the field only leaves on branches of ripe 8 × 4–5 mm. Adult leaves 5–10(–12) mm
mature trees, preferably from sun-exposed foliage, wide P. longifoliolatus
should be taken into account. Male (pollen) and 9a. Shrubs or small trees. Adult leaves 5–11 mm
female cones and seeds (they occur on dioecious wide. Pollen cones always solitary. Receptacles
trees) are described as mature. when ripe turning from red to blackish
P. pallidus 801
1a. Adult leaves 0.8–5 cm long 2 9b. Trees. Adult leaves (6–)8–15(–18) mm wide.
1b. Adult leaves (2.5–)4–14(–15) cm long 3 Pollen cones solitary or more often with 2–3 to-
2a. Trees. Adult leaves 2.5–5 cm long, 6–9 mm gether. Receptacles when ripe red 10
wide P. affinis 10a. Buds 2–5 mm long, sometimes with a few long-
2b. Shrubs. Adult leaves 0.8–2.2 cm long, 1.7–2.3 er outer scales to 8 mm. New leaves flushing
mm wide P. gnidioides red P. neriifolius
3a. Buds small (usually ca. 2 mm long); bud scales 10b. Buds 2–3 mm long, without longer outer scales.
imbricate, obtuse or short triangular 4 New leaves flushing pale whitish green
3b. Buds larger (usually more than 3 mm long); P. insularis
bud scales free, erect, spreading or outer bud
scales recurved, triangular-lanceolate, elongat-
Key to the species of Podocarpus in New
ed or acuminate 6
Zealand
4a. Shrubs. Adult leaves 3–5(–6) mm wide, with a
central groove on adaxial (upper) side When using this key in the field only leaves on
P. novae-caledoniae branches of mature trees, preferably from sun-­
4b. Trees. Adult leaves (5–)7–18 mm wide, with an exposed foliage, should be taken into account. Male
obtusely raised midrib on the adaxial side 5 (pollen) and female cones and seeds (they occur on
5a. Adult leaves (9–)11–18 mm wide. Seeds includ- dioecious trees) are described as mature.
ing the epimatium without a distal crest
P. lucienii 1a. Shrubs. Pollen cones 1–2.5 mm wide 2
5b. Adult leaves (5–)7–11(–13) mm wide. Seeds in- 1b. Trees. Pollen cones 3–8 mm wide 3
cluding the epimatium with a (faint) crest 2a. Adult leaves (0.5–)0.8–2 cm long, 1.4–2 mm
P. sylvestris wide, acicular, pungent. Receptacles when ripe
6a. Low, decumbent shrubs. Bud scales free, erect, 5–7 mm long P. acutifolius
not spreading. Adult leaves oblanceolate (wid- 2b. Adult leaves 0.3–1 cm long, 2–2.5 mm wide, na-
est beyond the middle) P. decumbens viculate (boat-shaped) to ovate-linear, obtuse
6b. Erect shrubs or more often trees. Bud scales or mucronate. Receptacles when ripe 8–10 mm
free, spreading to recurved or reflexed. Adult long P. nivalis
leaves linear-lanceolate (widest at or below the 3a. Pollen cones solitary or more often in clusters
middle) 7 of 2–5, 20–30 × 5–8 mm. Adult leaves (1.5–)2–
7a. Buds with distinctly elongated outer scales 4–15 3(3.5) cm long, 3–5(–6.5) mm wide, with a nar-
mm long. Midrib on the adaxial (upper) side of row but conspicuous groove on adaxial (upper)
leaves obtusely raised. Seeds including the epi- side P. cunninghamii
matium 7–9 mm long 8 3b. Pollen cones solitary, sometimes in pairs, 12–
7b. Buds with triangular or lanceolate, sometimes 15(–20) × 3–4 mm. Adult leaves (0.5–)1–2.5(–3)
acuminate outer scales 2–6(–8) mm long. Mid- cm long, (1.5–)2–3.5(–4.5) mm wide, with a
rib on adaxial side of leaves acutely raised. Seeds faint groove on the adaxial side P. totara
including epimatium (8–)10–15 mm long 9
 7b. Receptacles when ripe 8–10 mm long. Adult
Key to the species of Podocarpus in Mexico,
leaves 1.5–4 cm long, spinescent P. buchii
Central America, and the Caribbean Islands
8a. Adult leaves (3–)7–15(–20) cm long, (12–)15–20
When using this key in the field only leaves on mm wide (juvenile leaves often substantially
branches of mature trees, preferably from sun-­ larger). Seeds including the epimatium 9–11
exposed foliage, should be taken into account. Male mm long P. magnifolius
(pollen) and female cones and seeds (they occur on 8b. Adult leaves (1.5–)3.5–9 cm long, (5–)6–14 mm
dioecious trees) are described as mature. wide. Seeds including the epimatium 7–8 mm
long 9
1a. Adult leaves (1–)1.5–3(–3.5) cm long, 3–5 mm 9a. Midvein on adaxial (upper) side of leaves form-
802 wide 2 ing a groove from base to apex 10
1b. Adult leaves (1.5–)2–15(–20) cm long, (3–)5– 9b. Midvein (midrib) on adaxial side of leaves
20(–22) mm wide 3 prominently raised, sometimes fading towards
2a. Adult leaves narrowly lanceolate (widest below apex, but not grooved 11
the middle), with apiculate-pungent or acute 10a. Adult leaves elliptic-linear (widest at the mid-
apex. Small trees to 16 m tall (often only 5–10 dle). Bud scales all imbricate P. hispaniolensis
m) P. urbanii 10b. Adult leaves lanceolate to linear-lanceolate
2b. Adult leaves oblanceolate to linear-oblanceo- (widest below the middle). Bud scales imbri-
late (widest above the middle), with acuminate cate but some outer bud scales may be spread-
apex. Shrubs or small trees to 8 m tall ing P. oleifolius
P. ekmanii 11a. Midrib on adaxial (upper) side of leaves slightly
3a. Buds (on leading shoots) at least 5 mm long, raised, fading towards the acute to apiculate,
often twice as long, with ovate-apiculate to sometimes pungent apex P. purdieanus
long acuminate, more or less spreading (outer) 11b. Midrib on adaxial side of leaves prominently
scales 4 raised, continuous to the acute or obtuse (or
3b. Buds (on leading shoots) not longer than 5 mm, weakly apiculate) apex 12
with ovate to triangular, rounded to apiculate 12a. Bud scales all imbricate. Adult leaves 4–10 cm
or short acuminate, mostly imbricate scales 6 long, lanceolate to linear-lanceolate (widest be-
4a. Adult leaves (7–)10–18(–22) mm wide, lanceo- low the middle) P. guatemalensis
late, with a grooved midvein on adaxial (upper) 12b. Bud scales free at apex. Adult leaves (2.5–)4–
side P. costaricensis 6(–7) cm long, elliptic to oblanceolate (widest
4b. Adult leaves 7–14 mm wide, linear-lanceolate to at or above the middle) P. trinitensis
linear or narrowly lanceolate, with a raised but
fading midrib (midvein) on adaxial side 5
Key to the species of Podocarpus in South
5a. Pollen cones solitary or in pairs, 50–60 × 3–4
America
mm. Bud scales ovate-apiculate, free at apex
P. coriaceus When using this key in the field only leaves on
5b. Pollen cones always solitary, 30–45 × 4–5 mm. branches of mature trees, preferably from sun-­
Bud scales long acuminate, more or less spread- exposed foliage, should be taken into account. Male
ing P. matudae (pollen) and female cones and seeds (they occur on
6a. Adult leaves 1.5–5.5 cm long, with an acute- dioecious trees) are described as mature.
pungent or spinescent apex. Buds small, with
ovate or rounded scales 7 1a. Buds longer than wide at base, with outer scales
6b. Adult leaves (1.5–)3.5–15(–20) cm long, with 9–15 mm long, acuminate or cuspidate, some-
an acute, acuminate or obtuse apex (if pun- times elongated to become more or less leaf-
gent, adult leaves longer than 6 cm). Buds up to like 2
5 mm long, with triangular or acuminate, rarely 1b. Buds (usually) not longer than wide at base,
ovate or rounded scales 8 with outer scales up to 6 mm long, of various
7a. Receptacles when ripe 10–15 mm long. Adult shapes but with a short apex and not elongated
leaves 2–5.5 cm long, acute-pungent 7
P. angustifolius
2a. Adult leaves (1.5–)2–5 cm long; midvein on Trees to 20 m tall but may be dwarfed at the
adaxial (upper) side of leaves a continuous highest altitudes P. sprucei
groove 3 11a. Bud scales acute, acuminate or cuspidate, erect
2b. Adult leaves (2–)4–13 cm long; midrib (mid- and free at apex 12
vein) slightly raised or obtusely raised, with or 11b. Bud scales broadly triangular, ovate or round-
without a medial groove towards leaf apex 5 ed, imbricate or rarely some outer scales free at
3a. Adult leaves linear-lanceolate, 2–4(–5) mm apex 14
wide, strongly spinescent (with a hard, spine- 12a. Adult leaves 4.5–7.5 cm long, 8–15 mm wide.
like apex) P. glomeratus Trees to 30 m tall P. celatus
3b. Adult leaves elliptic, linear-lanceolate or oblan- 12b. Adult leaves 2–4 cm long, 4–8 mm wide. Shrubs
ceolate, 4–8(–10) mm wide, acute or obtuse 4 or small trees to 10 m tall 13 803
4a. Adult leaves oblanceolate (widest above the 13a. Adult leaves ovate to ovate-linear, 7–8 mm wide
middle) with an obtuse apex. Outer bud scales (widest below the middle), apex acuminate
free but erect, the apices coming together P. acuminatus
P. aracensis 13b. Adult leaves oblanceolate to elliptic, 4–7 mm
4b. Adult leaves elliptic to linear-lanceolate (wid- wide (widest at or above the middle); apex
est at or below the middle) with an acute apex. acute or obtuse P. roraimae
Outer bud scales spreading P. rusbyi 14a. Midrib on adaxial (upper) side of leaves initial-
5a. Adult leaves (7–)8–13 cm long, with an obtusely ly raised but fading towards apex. Pollen cones
raised, not grooved midrib on adaxial (upper) long and very slender, 25–50 × 1.5–2 mm
side. Receptacles when ripe 8–12 mm long, dark P. salignus
purple P. salicifolius 14b. Midrib (midvein) continuously raised and/
5b. Adult leaves (2–)4–9(–11) cm long, with a or grooved, or a continuous groove not raised
slightly raised and grooved midrib on adaxial above general surface of leaf. Pollen cones short
side. Receptacles when ripe 6–8 mm long, red and slender, or longer than 20 mm but then at
or purple 6 least 3 mm wide 15
6a. Buds 3–4 mm long, with a few elongated out- 15a. Pollen cones solitary or more often 2–6 to-
er scales to 10 mm long. Leaf apices all acute. gether, pedunculate, 6–12 × 1.5–2 mm. Seeds
Seeds including the epimatium without an api- including the epimatium 4–5 mm long, glo-
cal crest P. sellowii bose, without an apical crest. Buds smaller than
6b. Buds 5–15 mm long, with ovate-cuspidate outer 3 mm P. transiens
scales all apically elongated. Leaf apices acute 15b. Pollen cones always solitary, sessile, more than
or obtuse. Seeds including the epimatium with 10 mm long. Seeds including the epimatium
a minute apical crest P. steyermarkii 6–11 mm long, not globose and with an apical
7a. Adult leaves (1.5–)2–4(–5) mm wide; apex an- crest (sometimes inconspicuous). Buds usually
gustate-acute or pungent 8 3–5 mm long 16
7b. Adult leaves (3.5–)4–15(–20) mm wide; apex 16a. Adult leaves oblanceolate or elliptic (widest at
acute or obtuse, sometimes acuminate 11 or above the middle), (1–)1.5–2.5(–3.5) cm long,
8a. Bud scales 4–6 mm long, apiculate or lanceo- 3.5–5 mm wide P. tepuiensis
late, outer scales erect or slightly spreading, al- 16b. Adult leaves lanceolate-linear to linear, some-
ways free at apex 9 times narrowly oblanceolate (usually widest
8b. Bud scales shorter than 4 mm, broadly triangu- below the middle), (3–)4–13(–20) cm long, (5–
lar, imbricate 10 )6–18(–20) mm wide 17
9a. Adult leaves 1.5–3 cm long P. nubigenus 17a. Adult leaves (12–)15–20 mm wide, highly vari-
9b. Adult leaves (2.5–)4–8(–9) cm long able in length but up to 20 cm or more long
P. parlatorei P. magnifolius
10a. Midrib (midvein) on adaxial (upper) side of 17b. Adult leaves (5–)6–14(–18) mm wide, some-
leaves continuously raised. Small trees to 12 m what less variable in length and not longer than
tall P. lambertii 12–13 cm 18
10b. Midvein on adaxial side of leaves a continuous 18a. Midrib on adaxial (upper) side of leaves sharply
groove, not raised above general surface of leaf. and continuously raised, not lying in a (­shallow)
 groove. Pollen cones 10–15 mm long 4a. Adult leaves 4–5 mm wide, with a nearly absent
P. guatemalensis (inconspicuous) midrib on adaxial (upper)
18b. Midrib on adaxial side of leaves obtusely raised, side P. lawrencei
lying in a groove or becoming grooved to- 4b. Adult leaves 2–2.5 mm wide, with a central
wards leaf apex. Pollen cones 20–30 mm long groove on adaxial side P. nivalis
19 5a. Bud scales lanceolate, to 6 mm long, erect or
19a. Adult leaves pendulous. Pollen cones 6–7 mm slightly spreading. Midrib on adaxial (upper)
wide P. pendulifolius side of leaves raised but fading towards apex.
19b. Adult leaves spreading. Pollen cones 3–4 mm Pollen cones 2–4 together, sessile. Receptacles
wide 20 when ripe 7–8 mm long; seeds including the
804 20a. Adult leaves with a narrow midrib in a me- epimatium 8–9 × 6–7 mm P. nubigenus
dial groove, not elevated above general adaxial 5b. Bud scales obtuse or acute, 2–3 mm long, imbri-
(upper) surface of leaf. Bud scales rounded or cate or a few with a free apex. Midvein (midrib)
ovate, all imbricate P. oleifolius on adaxial side of leaves a faint groove. Pollen
20b. Adult leaves with an obtusely raised or more cones solitary or sometimes in pairs, short pe-
or less flat and dorsally grooved midrib. Bud dunculate. Receptacles when ripe 4.5–5.5 mm
scales (broadly) triangular, with some outer long; seeds including the epimatium 3–3.5 ×
scales free at apex P. brasiliensis 2–2.5 mm P. totara
6a. Buds very small (< 3 mm), with imbricate
scales 7
Key to the species of Podocarpus commonly
6b. Buds larger, with outer scales longer than 3
in cultivation
mm and spreading or recurved (at least free at
The species of Podocarpus keyed out here can be apex) 8
found in cultivation within and without the areas in 7a. Adult leaves narrowly linear-lanceolate, (3–)4–
which they also occur naturally. If a species is only 7 mm wide. Pollen cones very slender, 25–50 ×
known to have been planted within the area (in the 1.5–2 mm. Receptacles when ripe 5–6 mm long;
sense of the keys) in which it also occurs naturally, it seeds including epimatium 7–8 × 4–5 mm
is here excluded because then it can be found in the P. salignus
key to the species of that area. Species that are only 7b. Adult leaves linear-lanceolate, (5–)7–14(–17)
known or assumed to be grown in a few botanic gar- mm wide. Pollen cones more robust, 10–20 ×
dens and/or arboreta are also not included here. In 2–3 mm. Receptacles when ripe 15–25(–30) mm
cultivation means here: planted in an amenity, for- long; seeds including the epimatium 15–20 ×
estry or horticultural context. When using this key 12–15 mm P. elatus
in the field only leaves on branches of mature trees, 8a. Adult leaves (3–)4–5(–7) mm wide; midrib on
preferably from sun-exposed foliage, should be tak- adaxial (upper) side of leaves fading towards
en into account. Male (pollen) and female cones and apex P. elongatus
seeds (they occur on dioecious trees) are described 8b. Adult leaves (3–)4–15(–18) mm wide; midrib
as mature. on abaxial side of leaves acutely raised and con-
tinuous (if indistinct or fading towards apex,
1a. Adult leaves 0.3–3 cm long, 1.4–5 mm wide 2 leaves wider than 6 mm) 9
1b. Adult leaves (1.5–)2.5–14(–15) cm long, (3–)4– 9a. Bud scales on leading shoots long acuminate,
15(–18) mm wide (if shorter than 3 cm, usually 8–12 mm P. matudae
wider than 4 mm) 6 9b. Bud scales on leading shoots not long acumi-
2a. Always shrubs with multiple stems 3 nate, 2–5 (rarely to 8) mm long 10
2b. Always trees with single stems 5 10a. Adult leaves (2–)3–5(–7) cm long. Bud scales
3a. Adult leaves acicular, 1.4–2 mm wide, pungent. erect, with free apices close together. Pollen
Bud scales acute with free apices P. acutifolius cones solitary, 8–25 × 6–8 mm P. costalis
3b. Adult leaves ovate-linear to linear, or naviculate 10b. Adult leaves (2–)3.5–12(–15) cm long. Bud
(boat-shaped), obtuse or mucronate. Bud scales scales spreading or recurved, rarely erect. Pol-
(broadly) triangular, imbricate 4 len cones solitary or more often 2–5 together,
15–40(–60?) × 2–4 mm 11
11a. Midrib on adaxial (upper) side of leaves indis-
Vernacular names
tinctly raised, leaf apices obtuse or mucronate.
Pollen cones solitary or in pairs. Small bracts No common names have been recorded for this spe-
(‘foliola’) basal to receptacles absent cies.
P. latifolius
11b. Midrib on adaxial side of leaves acutely raised.
Description
Leaf apices acute, acuminate or obtuse. Pollen
cones 2–5 together (sometimes solitary). Small Small trees 4–5 m tall; trunk to 20 cm d.b.h. Bark
bracts basal to receptacles present 12 not described. Branches forming a dense crown.
12a. Adult leaves from nearly oval (the shortest) to Foliage branchlets slender, short and ascending
linear (with parallel margins) in longer leaves to erect, terete, with fine grooves between the leaf 805
P. polystachyus bases, terminating in small, globose buds with erect,
12b. Adult leaves linear-lanceolate or lanceolate 4–6 mm long, cuspidate scales that are free at apex.
(widest below the middle), the shortest leaves Leaves ovate to ovate-linear, 3–4 cm long, 7–8 mm
sometimes oblanceolate (widest above the mid- wide, more or less abruptly narrowing to a short pet-
dle) 13 iolate or sessile base, terminating in an acuminate
13a. Bud scales caudate, with a narrow, recurved apex; leaf texture coriaceous; leaf colour deep green
apex. Shrubs or small trees. Small bracts (‘fo- above, dull green below. Midrib on adaxial (upper)
liola’) basal to receptacles also recurved side a continuous groove, on abaxial side obtusely
P. macrophyllus raised. Stomata very small, in two bands of numer-
13b. Bud scales lanceolate or triangular, spreading ous intermittent lines on abaxial side. Pollen cones
with a more or less straight apex. Potentially not observed. Seed cones axillary, solitary on a short
large trees. Small bracts basal to receptacles not peduncle; receptacles consisting of an axis with 3–4
recurved 14 partly fused bracts, one or two of which are smaller
14a. Adult leaves (4–)6–10 mm wide. Pollen cones and sterile and occur below the usual (second) ster-
15–25(–30) × 2–2.5 mm. Receptacles when ripe ile bract, the whole swelling to ca. 6 mm long and
9–16 mm long; seeds including the epimatium 3–4 mm wide and green, becoming red or purple at
ca. 9 mm long, green or glaucous P. fasciculus maturity. Seeds solitary, sometimes 2 per receptacle,
14b. Adult leaves (5–)6–15(–18) mm wide. Pollen obliquely ovoid, including the epimatium ca. 7 mm
cones 20–40(–60?) × 2–3.5 mm. Receptacles long and 4 mm wide, with a prominent distal crest,
when ripe 7–12 mm long; seeds including the green. Seed proper not observed.
epimatium (8–)10–15 mm long, purplish green
or blackish purple 15
Distribution
15a. Receptacles subtended by two 2–6 mm long
bracts (‘foliola’); epimatium around seeds with- Brazil: Amazonas (Serra da Neblina); Venezuela:
out a crest. New leaves flushing red P. neriifolius Amazonas (Sierra de la Neblina), Bolivar (Chi-
15b. Receptacles subtended by two minute, 1–2 mm mantá, SW Amuri-tepui).
long bracts; epimatium around seeds with a TDWG codes: 82 VEN 84 BZN-AM
crest. New leaves flushing yellowish green
P. subtropicalis Ecology
Podocarpus acuminatus occurs on upper slopes and
summits of some of the tepuis (table mountains)
Podocarpus acuminatus de Laub., Novon 2 (4): 329.
formed of sandstone in southern Venezuela and the
1992. Type: Brazil: Amazonas, Serra da Neblina,
border with Brazil. It is a sun-exposed shrubby tree
Pico da Neblina, C. Farney & B. F. E. Pesscoal 870
in closed dwarf forest, often growing among rocks,
(holotype MO).
at altitudes between 1900 m and 2400 m a.s.l.

Etymology Conservation
The species epithet refers to the apex of the leaves, Podocarpus acuminatus is known only from two
which tapers to a narrow point. widely disjunct localities in remote areas. The area
 of occupancy (AOO) is less than 500 km2 but the at anthesis; microsporophylls triangular, with two
remoteness of the localities and relative or perhaps globose pollen sacs. Seed cones axillary, solitary on
even completely undisturbed nature of the forests on ca. 5 mm long peduncles, consisting of an axis with
the tepuis and mountains makes it likely that no im- 2 obtuse bracts which fuse to become a swollen, suc-
mediate threats are present. However, road access to culent, red receptacle 5–7 mm long, subtending a
the Serra da Neblina area from Brazil has now been single, small, narrowly ovoid seed 4–5 mm long in-
established and, as experience shows, may pose a cluding the glaucous green epimatium. Seed proper
threat to the forests in the mountains. It is therefore not observed.
thought prudent to flag this rare species with NT.
IUCN: NT
Taxonomic notes
806
It is commonly stated that Thomas Kirk discovered
Uses
this species in the upper part of Buller Valley, Nel-
No uses have been recorded of this species. son, New Zealand (near the outflow of Lake ­Rotaiti),
but his collection from there is only one element
cited in the protologue, as he also acknowledges
Podocarpus acutifolius Kirk, Trans. & Proc. New T. F. Cheeseman for further specimens. At Kew (K)
Zealand Inst. 16: 370. 1884. Type: New Zealand: is a sheet with a mixture of male and sterile (female?)
South Island, Nelson, Buller Valley, near outflow of branchlets collected by Cheeseman at Lake Rotaiti.
Lake Rotaiti, T. Kirk s.n. (syntype WELT). Fig. 255 Since the plant is dioecious, these constitute more
than one collection and we do not know what Kirk
saw of this (if any, they are likely duplicates). A de-
Etymology
tailed investigation is needed to select a lectotype
The species epithet refers to the sharp pointed leaves. from these (and other) syntypes; preferably choos-
ing a Kirk specimen in a New Zealand herbarium.
Vernacular names
Distribution
Needle-leaved totara
New Zealand: South Island (Malboro Straits to S
Westland).
Description
TDWG codes: 51 NZS
Low decumbent or erect shrubs 50–150 cm tall, or
small, bushy trees to 9 m tall with a trunk to 40 cm
Ecology
diam. Branches numerous, on larger shrubs and
trees ascending, otherwise spreading, branching ir- Podocarpus acutifolius is a shrub or bushy tree oc-
regularly. Foliage branchlets lax and slender or more curring in lowland to montane forest and scrub.
rigid in small trees, finely grooved between leaf According to Allan in Flora of New Zealand 1: 108
bases, glabrous, light brown turning grey; terminal (1982) the taller growing tree form is possibly of
buds conical, 2–4 mm long, with free, acute scales. hybrid origin (other parent not stated, but likely
Leaves spirally arranged, spreading at wide angles P. ­cunninghamii).
from shoot, acicular, (5–)8–20 mm long, 1.4–2 mm
wide, coriaceous, rigid, straight or slightly curved, Conservation
flattened; narrowed base twisted and derurrent;
IUCN: LC
apex pungent. Midrib inconspicuous on both sides;
leaf leaf colour green to yellowish green or bronze
Uses
green above, similar but with whitish stomatal lines
below. Stomata on abaxial side in two bands of inter- Needle-leaved totara is rare in cultivation. Under
mittent lines separated by a flat midrib. Pollen cones garden conditions it sometimes grows into a small,
axillary on short peduncles, solitary or with 2–4 to- bushy and open-crowned tree to 5 m tall, but prov-
gether, subtended by 3–4 scarious, acuminate bracts, enances from low creeping shrubs may stay in that
slender cylindrical, 12–20 mm long, 2–2.5 mm wide habit. It should be a species suitable to be trained
and pruned into hedges, while the decumbent form TDWG codes: 60 FIJ
with its often bronze foliage is an attractive shrub
for rockeries.
Ecology
Podocarpus affinis has been found in rainforest on
Podocarpus affinis Seem., Fl. Vitiensis: 266. 1868.
or near the summits of a few mountains in Viti Levu,
Type: Fiji: Western Division, Viti Levu, Voma Peak,
where it is locally common growing in the under-
B. C. Seemann 574 (holotype K).
storey or dominant in the canopy of low forest. The
altitudinal range is 900–1200 m a.s.l.
Etymology
The species epithet means ‘allied to’; Seemann 807
Conservation
thought it was allied to Podocarpus elatus.
Doyle (1998) has listed this species as Endangered
Vernacular names (EN) on grounds of its very limited distribution and
apparent decline and fragmentation of its habitat. It
kuasi (Viti Levu)
was listed as Vulnerable (VU) by the same author in
the Global Redlist of Conifers (Farjon & Page, 1999)
Description
under the 1994 A-criterion which estimates decline
Small to medium size tree to 15–20 m tall. Bark not only. Since these criteria have changed in 2001, a re-
described. Branches short and spreading; foliage assessment of this species seems necessary.
branches towards ends of main branches slender, IUCN: VU (A1c + 2c)
terete, terminating in small buds with more or less
erect, free, narrowly triangular, acute scales. Leaves
Uses
on seedlings similar to those on mature trees, 2.5–5
cm long, oblanceolate to elliptic or linear-elliptic, The wood of this species, according to Seemann, was
6–9 mm wide, widest at or above the middle, co- used for outrigger canoes by “the natives”; presum-
riaceous, gradually tapering to a petiolate base; ably it was used for other purposes as well. Deple-
apex obtuse to rounded; margins slightly revolute, tion of the resource has greatly diminished this use
lustrous green above, dull brownish green below. at present. The species is not known in cultivation.
Midrib narrow (ca. 0.5 mm wide) and acutely raised
on adaxial (upper) side, similar but fading towards
apex on abaxial side. Stomata very small, in numer- Podocarpus angustifolius Griseb., Cat. Pl. Cubensis:
ous irregular and intermittent lines on abaxial side. 217. 1866. Type: W Cuba, [“Cuba occidentalis”],
Pollen cones and seed cones remain unknown. C. Wright 3188 (holotype not located, isotypes BM,
K).
Taxonomic notes
Podocarpus aristulatus Parl., in Candolle, Prodr. 16
Although the leaves of this species are distinct (but (2): 513. 1868; Podocarpus angustifolius Griseb. var.
reminiscent of P. pilgeri) at least among the podo- aristulatus (Parl.) Staszk., Fragm. Fl. Geobot. 33: 77.
carps of Fiji and New Caledonia, to date no speci- 1988.
mens have been collected with either pollen or seed Podocarpus leonii Carabia, Caribbean Forest. 2: 92.
cones. Apparently, this tree rarely bears these repro- 1941. Podocarpus angustifolius Griseb. var. leonii
ductive organs and all herbarium specimens cited by (Carabia) Staszk., Fragm. Fl. Geobot. 33: 77. 1988,
previous authors, as well as seen in the Kew Herbari- [“leonis”].
um (K) are sterile. Acceptance of this taxon remains Podocarpus victorinianus Carabia, Caribbean For-
therefore more or less provisional. est. 2: 92. 1941 (nom. illeg., Art. 52); Podocarpus
angustifolius Griseb. var. leonii (Carabia) Staszk. f.
victorinianus (Carabia) Staszk., Fragm. Fl. Geobot.
Distribution
33: 77. 1988.
Southwestern Pacific: Fiji Islands (Viti Levu, Namo-
si, Voma Peak).
 (P. ­aristulatus) and Wright 3188 (P. angustifolius)
Etymology
gives a difference in maximum width of leaves of
The species epithet refers to the narrow leaves. 3 mm (8 mm resp. 5 mm). The specimen at BM of
Wright 1461 has leaves to 12 mm wide and none nar-
rower than 8 or 9 mm. Because Wright 1461 at K is
Vernacular names
a male specimen and the one at BM female, they
No common names have been recorded for this spe- cannot have been taken from the same tree, which
cies. means that Wright 1461 is a mixed collection. Cara-
bia (op. cit.) described a new species P. victorinianus
based on Wright 1461 at GH. His observation that it
Description
808 has 1–1.2 cm wide leaves is in agreement with Wright
Large shrubs or small trees to 12 m tall; trunk to 1461 at BM, which, like the specimen he cites from
50 cm d.b.h. Bark thin, smooth, brown weathering GH, is female. Gray & Buchholz (1948) described
grey. Branches on young trees subverticillate and the two species on the same page of their revision
spreading, ascending on multi-stemmed shrubs. Fo- of American podocarps, but did not indicate any
liage branches slender, terete, with fine longitudinal difference other than the width of leaves. Parlatore
grooves between the leaf bases, terminating in small, (1868) cited Wright 1461 and Wright 3788 (typo-
ovoid or subglobose buds with imbricate, ovate, car- graphic error for Wright 3188) under P. aristulatus
inate outer scales with scarious margins terminating [“aristulata”] and wrote of the latter specimen that
in an acuminate apex. Leaves on saplings and ma- it has 4–5 mm wide leaves. Florin (1934) considered
ture trees similar, more or less remotely spaced espe- them synonyms. Other specimens at K from Cuba
cially on vigorous shoots, patent, narrowly elliptical appear to fall more or less between leaf width meas-
(shortest and widest leaves) or lanceolate-linear urements of 5–8 mm and are difficult to assign to
to linear, 2–5.5 cm long, (3–)5–10(–12) mm wide, either P. angustifolius or P. aristulatus.
straight or curved downward, gradually narrowing
to a short petiolate base and to an acute-pungent or
Distribution
spinaceous apex; margins revolute; leaf colour lus-
trous dark green above, dull green below. Midrib on Caribbean: Cuba (Granma, Holguín, Pinar del Rio,
adaxial (upper) side narrow, slightly raised but often Sancti Spiritus).
fading towards apex, acutely raised on abaxial side TDWG codes: 81 CUB
and continuous almost to apex. Stomata very small,
in intermittent lines on either side of the abaxial
Ecology
midrib. Pollen cones axillary, solitary, sessile, cy-
lindrical, elongating to 15–20 mm, 2.5–3 mm wide Podocarpus angustifolius occurs in montane rain-
at anthesis; microsporophylls broadly ovate, obtuse forests of Cuba, often on laterite soils derived from
with minutely erose-denticulate margin, bearing serpentine. Its altitudinal range is undetermined, but
two basal ovoid-oblong pollen sacs. Seed cones ax- several herbarium collections were made around
illary, short pedunculate; receptacles with 2–3 un- 800 m a.s.l. It is associated with mostly angiosperm
equal, fused bracts, swelling to 10–15 mm long and shrubs and trees, such as Magnolia cubensis, Cojoba
7–10 mm wide, succulent and ripening to bright arborea, Beilschmiedia pendula, Octea spp., Lauro­
red. Seeds solitary, including the epimatium 7–10 cerasus occidentalis, Mataybe domingensis, etc., and
mm long, narrowly ovoid with a small, 1–2 mm long can occur in low forest canopy as well as in more
crest, green. Seed proper ovoid, smooth, dark yel- open, sun-exposed vegetation on rocky sites. In
lowish brown (‘tawny’). some localities it has been found to be abundant,
with regeneration representing all age classes in un-
disturbed forest, but nearly absent in disturbed areas.
Taxonomic notes
The distinction between Podocarpus aristulatus and
Conservation
P. angustifolius appears to be based on the (maxi-
mum) width of leaves. A comparison between This species, now known not only from western
the type collections (specimens at K) Wright 1461 Cuba but also from central and eastern parts of the
island, is nevertheless uncommon, although locally narrowing to a 2–5 mm long petiolate base, more
abundant. Its total area of occupancy (AOO) is less abruptly ending in an acute (juvenile plants) or ob-
than 500 km2 and populations are widely separated tuse apex. Margins revolute, leaf texture coriaceous;
and/or fragmented. Although not usually a timber midrib on adaxial (upper) side a continuous narrow
tree, it is declining mainly because of forest clear- groove, on abaxial side raised, or nearly flat in sicco.
ance to make way for agriculture. It regenerates Stomata on abaxial side in numerous intermittent
poorly or not at all in disturbed areas, even if these lines on either side of midrib. Pollen cones axillary,
are left for secondary forest regeneration, probably solitary or in clusters of 2–3, with a scaly 7–9 mm
because of low competitiveness. long peduncle, cylindrical, 17–20 mm long, 2–2.5
IUCN: EN (B1+2c) mm wide; microsporophylls triangular, with two
globose pollen sacs. Seed cones not observed. 809
Uses
Taxonomic notes
No commercial uses have been recorded of this spe-
cies. Larger trunks of trees growing in forest may This species remains poorly known due to insuffi-
have been logged and used for timber, but probably cient collecting. Some additional herbarium speci-
only locally. It is locally planted in Cuba as an orna- mens, collected from the Cerro de la Neblina in
mental shrub or small tree. Venezuela, were mentioned in the protologue (De
Laubenfels & Silba, 1988) but these are doubtfully
identified as having characters shared with P. ara­
Podocarpus aracensis de Laub. & Silba, Phytologia censis as well as P. buchholzii, another of David de
65: 330. 1988. Type: Brazil: Amazonas, Serra Laubenfels’s poorly known species from Venezuela.
Araca, [“margins of R. Serra Araca”], N. A. Rosa & An additional herbarium collection (O. Huber 11873,
S. B. Lira 2317 (holotype MG). K), not cited in the protologue, but identified by
De Laubenfels in 1992 as belonging to Podocarpus
aracensis, was collected in 1986 on Cerro Yaví in
Etymology
­Amazonas Territory, Venezuela, nearly 500 km to
This species was named for the mountain, Serra the north of Cerro de la Neblina and 550 km NNW
Araca, where it was first found. of Serra Araca, the type locality in Brazil.

Vernacular names Distribution

No common names are known for this species. Brazil: Amazonas (Serra Araca, Cerro Neblina?);
Venezuela: Amazonas (Cerro Yaví).
TDWG codes: 82 VEN 84 BZN-AM
Description
Shrubs or small trees 2–6 m tall; trunk diam. 10–20
Ecology
cm or more. Branches spreading, in older plants
forming a dense crown. Foliage branchlets slender, This species has been found on sandstone massifs
terete, finely grooved between leaf bases, terminat- along streams in shrubby vegetation or dwarf for-
ing in conical buds with erect, elongated outer scales est at altidudes around 1200 m a.s.l. in Brazil and at
4–9 mm long, ca. 2 mm wide at base, constricted 2100 m in Venezuela.
to a 0.5 mm wide distal part. Leaves on saplings
and juvenile (or shaded?) plants remotely placed
Conservation
along shoots, on older shrubs and trees and on sun-­
exposed branches crowded at the tips, spreading to IUCN: LC
nearly erect, lanceolate-linear on juvenile plants,
oblanceolate (widest above the middle) on mature
Uses
shrubs and trees, 5–8 cm long on juvenile plants,
2–4.5 cm long on mature shrubs and trees, 4–7(–9) No uses are known of this species, which occurs re-
mm wide (widest on juvenile plants), gradually mote from human habitation.
 Podocarpus archboldii N. E. Gray, J. Arnold Arbor. Seed cones axillary, solitary on a 5–10 mm long pe-
39: 452. 1958. Margbensonia archboldii (N. E. Gray) duncle; receptacles with 2 minute bracts at base and
A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc. often two protruding lateral bracts, obliquely bilo-
Isp. Prir., Otd. Biol. 103 (1): 59. 1998. Type: Indone- bate, ca. 10 mm long, swelling with a truncate distal
sia: Papua, Idenburgh River, Bernhard Camp, 4 km end, red or purple-black and succulent when ripe.
SW of camp, L. J. Brass 13121 (holotype A). Seeds enclosed in a smooth epimatium, light green
when ripe, ovoid, 8–9 × 5–6 mm, with a distal crest.
Seed proper not observed.
Etymology
The species epithet commemorates Richard Arch-
Distribution
810 bold, who sponsored the botanical expedition to
New Guinea on which this species was first found. New Guinea.
TDWG codes: 43 NWG-IJ NWG-PN
Vernacular names
Ecology
mu, soa (Papua), sarau (PNG); probably many other
names in local languages. Podocarpus archboldii is a canopy tree occurring
scattered or locally common in evergreen tropical
montane forest. The altitudinal range is (720–)1500
Description
m to 2600 m a.s.l. The largest trees occur in closed
Trees to 45 m tall, to 100 cm d.b.h., bole erect, forests dominated by Castanopsis spp. with a canopy
straight, sometimes fluted at base. Bark smooth, to 50 m tall; trees become smaller and stunted in
thin, longitudinally and often more or less spirally mossy forest which often occurs on mountain ridge
fissured, brown weathering grey; inner bark cream, tops over rocky terrain. Here soils are less well devel-
fibrous. Branches spreading, forming a narrow or oped and other conifers, e.g. Agathis labillardierei,
rounded crown. Foliage branchlets terete, stout, Dacrydium novoguineense, Falcatifolium ­papuanum,
more or less grooved, glabrous, terminating in 3–5 ­Papuacedrus papuana, and ­Retrophyllum vitiense,
mm long, nearly globular buds with slightly spread- may join Podocarpus archboldii, while angiosperms
ing, triangular to lanceolate outer scales and imbri- become less dominant.
cate, rounded inner scales. Leaves on juvenile plants
short petiolate, linear-lanceolate, (3–)6–14(–18) cm
Conservation
long, 10–16 mm wide, straight or slightly curved,
gradually narrowed at base, tapering to an acute IUCN: LC
apex. Leaves of mature trees usually shorter, linear-
lanceolate, (3–)5–10(–12) cm long, 8–13 mm wide,
Uses
straight or slightly curved, coriaceous, sometimes
convex or with revolute margins, short petiolate at a Podocarpus archboldii is a valuable timber tree
gradually or abruptly narrowing base, tapering to an where it attains large sizes with a clear, straight bole.
acute or obtuse apex. Midrib obtusely raised on both Its wood is used as roundwood for masts, spars and
sides, usually narrower on adaxial side; leaf colour poles, in house construction as beams, in high-
dark green above, pale green below, new leaves grade construction for flooring, joinery and other
flushing light yellowish green. Stomata very small, carpentry, for furniture and cabinet work, veneer, to
in numerous irregular lines on abaxial (under) side make boxes, and for match sticks. Its traditional uses
on either side of midrib. Pollen cones axillary, soli- include village house construction, household uten-
tary, sessile or short pedunculate with several large sils and wood carving. This species has been found
basal bract scales, cylindrical, elongating to 30–40 either spared from the forest or planted in village
mm, 6–8 mm wide; microsporophylls spreading, dancing grounds.
apical part elongated, with two globose pollen sacs.
Podocarpus atjehensis (Wasscher) de Laub. ex × 7–8 mm, whitish or bluish pruinose, without a
Silba, Phytologia Mem. 7: 60. 1984. Podocarpus crest. Seed proper not observed.
neriifolius D. Don var. atjehensis Wasscher, Blumea
4 (3): 450. 1941; Margbensonia atjehense (Wasscher)
Taxonomic notes
A. V. Bobrov & Melikyan, Bjull. Moskovsk.
Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998. Type: Formerly known as Podocarpus neriifolius D. Don
Indonesia: Sumatera, Aceh, Gunung Kemiri, var. atjehensis Wasscher, this taxon was elevated to
C. G. G. J. van Steenis 9614 (holotype L). species rank by David. J. de Laubenfels (1985), who
classified the two species in different sections based
on vegetative characters. Although De Laubenfels
Etymology
should be credited for the taxonomy, it was John Silba 811
The species epithet refers to Atjeh [now to be spelled (op. cit.) who first validly published the new combi-
Aceh], the semi-autonomous region of northern Su- nation, citing the full reference to the basionym.
matera in Indonesia.
Distribution
Vernacular names
Malesia: Sumatera (Aceh, Gajo Lands); Papuasia:
No common names have been recorded for this spe- New Guinea (Papua, Wissel Lakes).
cies. TDWG codes: 42 SUM 43 NWG-IJ

Description Ecology
Small trees 8–15 m tall, to 25 cm d.b.h. Bark not de- Podocarpus atjehensis was discovered and described
scribed. Branches spreading and ascending, forming (as a variety of P. neriifolius) from the summit area
a pyramidal to rounded crown. Foliage branchlets of a high mountain in northern Sumatera growing
terete, with narrow ridges from decurrent leaf peti- in mossy dwarf forest and shrubland. The type col-
oles; leading branches terminating in large buds lection was gathered at 2900 m a.s.l. on a slope be-
6–14 mm long, 5–8 mm wide, with broadly triangu- low the summit of Gunung Kemiri, and it may (or
lar scales, the outer scales with apiculate, recurved may not) occur near or on the summit of nearby
tips, the inner scales shorter and more or less im- Gunung Leuser, the highest mountain in the area at
bricate. Leaves of young plants linear, usually de- 3145 m. The other locality reported for this taxon is
flected, to 18 cm long and 9 mm wide; leaves of adult in New Guinea at 1800 m a.s.l.
trees linear-lanceolate, petiolate at base, strongly
deflected, 6–11 cm long, 5–8 mm wide, coriaceous,
Conservation
gradually narrowing at both ends; apex long acute;
midrib on adaxial (upper) side less than 0.4 mm This species is only known from two very disjunct
wide and acutely raised, on abaxial (lower) side ca. locations. This type of disjunction is also reported
1 mm wide and raised to a prominent flat ridge; leaf from other groups of organisms. In both locations
colour dark green above, light green below, flushing there is a considerable human footprint on the land-
pink or rosa. Pollen cones axillary, solitary, sessile, scape and although we do not know to what extent
subtended by broadly ovate-triangular bud scales, this affects this species, it is assumed that some de-
long cylindrical, 25–30 mm long, 4–4.5 mm wide at cline has occurred. IUCN’s Conifer Specialist Group
anthesis; microsporophylls with a small triangular therefore wish to flag this species as Near Threat-
apex and two large pollen sacs. Seed cones axillary, ened.
solitary, on a 8–15 mm long peduncle; receptacles IUCN: NT
with one (or two?) 2–4 mm long basal bract(s) (‘fo-
liola’), unequally bilobate with two bract tips, swell-
Uses
ing to 9–11 × 6–7 mm, succulent and red at maturity.
Seeds including the epimatium ovoid-globose, 9–11 No uses have been recorded of this poorly known
species.
 Podocarpus borneensis de Laub., Blumea 30
Distribution
(2): 266. 1985. Type: Malaysia: Sabah, Sandakan
District, Bukit Tawai, D. J. de Laubenfels P 702 Indonesia: Kalimantan (including Karimata Island);
(holotype L). Malaysia: Sabah, Sarawak.
TDWG codes: 42 BOR-KA BOR-SB BOR-SR
Podocarpus polystachyus R. Br. ex Endl. var. rigidus
Wasscher, Blumea 4 (3): 460. 1941.
Ecology
Podocarpus borneensis occurs in mossy forest on
Etymology
mountain ridges, often in rocky, exposed sites as a
812 The species epithet refers to the island of Borneo, small or stunted tree. Downslope, where the forest
where it is endemic. becomes taller and has a closed canopy, scattered
trees may reach into the canopy and attain 20 m
or more. It also occurs in ‘keranga’ forest on nearly
Vernacular names
white sand and it was once or twice collected from
bisit, bubung, buloh (local names in Sarawak) swamp forest at low altitude (ca. 350 m) on peaty soil.
Its more common altitudinal range is from 700 m to
2100 m a.s.l. On the Meurong Plateau (sandstone),
Description
it may become the dominant tree in some localities,
Small to medium sized trees 5–10(–23) m tall, up to elsewhere it is mixed with various conifers (Agathis,
30 cm d.b.h. Bark not described. Branches spread- Dacrydium, Dacrycarpus, Podocarpus, Phyllocladus)
ing, forming a rounded or domed crown. Foliage and angiosperm shrubs and trees.
branchlets terete, more or less grooved, glabrous,
terminating in 4–10 mm long, 2–3 mm wide buds Conservation
with erect or slightly spreading, lanceolate to linear
IUCN: LC
outer scales. Leaves on juvenile plants short petio-
late, linear, to 16 cm long and to 14 mm wide, straight
Uses
or slightly curved, gradually narrowed at base, acute
or obtuse. Leaves of mature trees often crowded at No economic uses have been recorded of this spe-
the distal end of branchlets, spreading or ascending, cies. Its ecology and commonly low stature make
linear-ovate or elliptic, 3–7(–9) cm long, 8–13 mm it an unlikely target of timber logging. Where it is
wide, straight or slightly curved, short petiolate at a abundant, like on the Merurong Plateau, its wood
gradually narrowing base, tapering to an obtuse or is locally used for construction of houses and other
rounded (rarely acute) apex, thick coriaceous. Mid- carpentry work. It is not known to be in cultivation.
rib acutely raised on adaxial side, nearly flat on abax-
ial side, drying to a groove; leaf colour dark green
above, pale green below. Stomata very small, in nu- Podocarpus bracteatus Blume, Enum. Pl. Javae
merous irregular lines on abaxial (under) side on ei- 1: 88. 1827. Podocarpus neriifolius D. Don var.
ther side of midrib. Pollen cones axillary, solitary or bracteatus (Blume) Wasscher, Blumea 4 (3): 449.
in clusters of 2–3, sessile or short pedunculate with 1941. Type: Indonesia: Jawa, [locality not stated],
several basal bract scales, cylindrical, elongating C. L. Blume s.n. (holotype L).
to 35–50 mm, 2.5–3.5 mm wide; microsporophylls
triangular, spreading, with two pollen sacs. Seed
Etymology
cones axillary, solitary on a 2–3 mm long peduncle;
receptacles with 2 bracts (‘foliola’) 1–2 mm long at The species epithet describes the two conspicuous
base, obliquely bilobate, 6 mm long, swelling with a bracts at the base of the receptacle.
truncate distal end, green turning red and succulent
when ripe. Seeds including the epimatium ovoid, 6
Vernacular names
× 5 mm, the distal end cristate, green turning purple
or dark brown when ripe. Seed proper not observed. ki marak, ki pantjar, ki putri (Jawa); kayu unung un-
ung (Sumatera)
 tus. However, in the widespread species P. neriifolius
Description
basal bracts or ‘foliola’ as long as 5 or even 6 mm oc-
Trees to 40 m tall, to 1 m d.b.h., bole straight, in large casionally occur, and consequently this is a spurious
trees slightly fluted or rarely buttressed at base. Bark character. To base divisions in the genus on this sin-
smooth, thin, on large boles with narrow, longitudi- gle feature amounts to ‘artificial’ classification (see
nal flakes or strips, light brown weathering grey; in- taxonomic notes above under the genus).
ner bark pinkish or reddish brown, fibrous. Branches
spreading, forming a rounded or domed crown. Foli-
Distribution
age branchlets terete, more or less grooved, glabrous,
terminating in 5–12 mm long buds with spreading, Indonesia: Flores, Jawa, N Sumatera.
triangular to lanceolate outer scales. Leaves on ju- TDWG codes: 42 JAW LSI-LS SUM 813
venile plants short petiolate, linear-lanceolate, 15–23
cm long, 15–20 mm wide. Leaves of mature trees
Ecology
shorter and narrower, mostly linear-lanceolate or
lanceolate, (6–)8–12(–14) cm long, 9–14 mm wide, Podocarpus bracteatus is a scattered canopy tree of
straight or slightly falcate, gradually narrowing to a evergreen montane tropical rainforest, most com-
petiolate base, gradually tapering to an acute apex. monly found on forested volcanoes of Jawa at al-
Midrib acutely raised on adaxial (upper) side and titudes between 1000 m and 2600 m a.s.l. It has
0.4 mm wide, flat or obtusely (sometimes acutely) occasionally been collected from lower altitudes be-
raised abaxially; leaf colour dark green above, pale tween 400 m and 1000 m a.s.l. Its ecology is similar
green below, new leaves flushing red. Stomata very to that of P. neriifolius.
small, in numerous irregular lines on abaxial (un-
der) side on either side of midrib. Pollen cones axil-
Conservation
lary, solitary or in clusters of 2–3, sessile with several
basal, large, ovate-triangular, carinate bud scales, cy- Logging of larger trees of this species probably
lindrical, elongating to 25–50(–60) mm, 3.5–4 mm has had an impact on its conservation status, but
wide; microsporophylls spirally arranged, triangu- it seems too widespread and still reasonably com-
lar, spreading, with two globose pollen sacs. Seed mon at least in Jawa. If the botanical distinction with
cones axillary, solitary on a 10–20 mm long pedun- P. neriifolius is slight and not often made by foresters
cle; receptacles with 2 subulate bracts (‘foliola’) 4–5 in the field, it will be very difficult to know whether
mm long at base, swelling to become ellipsoid with a this species is declining or not. It is at present not
truncate distal end, 10–14 mm long, yellow turning considered to be threatened with extinction.
orange-red or bright red and succulent when ripe. IUCN: LC
Seeds at truncate end of the receptacle, enclosed in
a smooth epimatium, green turning purplish green
Uses
when ripe, ovoid-oblong or ovoid, 10–14 mm long,
ca. 7 mm wide. Seed proper ovoid, 7–10 mm long, This species is an important timber tree in Jawa. Its
slightly flattened. excellent wood is used for house construction and
carpentry and for making oars, spars and masts of
sailing vessels. More specialized uses requiring high
Taxonomic notes
grade timber are veneer, furniture making, cabinet
Podocarpus bracteatus is quite similar to P. neriifo­ making, interior trim, household utensils, and wood
lius, of which Wasscher (op. cit.) considered it to be carving. As far as known it is not in cultivation.
a mere variety. However, De Laubenfels (1985, 1988)
not only kept it as a distinct species, but classified
it in a different section: Longifoliolatus as opposed Podocarpus brasiliensis de Laub., Fl. Venezuela 11
to Foliolatus. Both are named for the ‘foliola’, i.e. the (2): 31. 1982. Type: Brazil: Distrito Federal, Brasilia,
two basal, subulate bracts below the receptacle. De Horto do Guará, E. P. Heringer 8034 (holotype NY).
Laubenfels held that these were about 2 mm long in
species of sect. Foliolatus, which contains P. nerii­ Podocarpus barretoi de Laub. & Silba, Phytologia 68:
folius, and “over 3 mm” in those of sect. Longifoliola­ 65. 1990.
 TDWG codes: 82 VEN 84 BZC-DF BZC-GO BZE-BA
Etymology
BZL-MG BZN-RO BZN-RM
The species epithet means ‘from Brazil’, where the
type specimen was collected.
Ecology
Podocarpus brasiliensis is apparently widespread
Vernacular names
and occurs both north and south of the Amazon Ba-
No common names have been recorded for this spe- sin (which is devoid of conifers in natural habitats).
cies. In Venezuela it is known from the tepuis, high and
isolated sandstone plateaux and table mountains,
814 where it is a component of dwarf forest above sand-
Description
stone bluffs, or occupies the transition zone to lower
Small trees to 15 m tall; trunk to 30 cm d.b.h. Bark altitude, taller evergreen rainforest. The altitudinal
not described. Branches spreading, forming a broad range is imperfectly known; where stated specimens
pyramidal to rounded crown. Foliage branchlets were collected between 800 m and 1600 m a.s.l. In
slender or stout, terete, with fine grooves between Brazil it occurs in the ‘cerrado’ which is mostly low,
the leaf bases; terminating in subglobose buds 3–4 open, and bushy vegetation, but here it is limited to
mm wide at base with (broadly) triangular scales, narrow strips of ‘gallery forest’ along streams, where
the inner ones imbricate and obtuse with scari- ground water is more reliably available. In the Mato
ous margins, the outer scales to 4 mm long with Grosso of Brazil a herbarium collection of this spe-
an acute or acuminate and erect, free apex. Leaves cies was made in “high forest” but this habitat was
on mature trees elliptic-linear to narrowly oblan- not further characterized.
ceolate, (3–)5–10(–13) cm long, (6–)9–14(–18) mm
wide, mostly straight, gradually tapering to a peti- Conservation
olate base, gradually or more abrubtly narrowing
IUCN: LC
to an acute or obtuse or slightly acuminate apex;
margins flat or weakly revolute; texture coriaceous;
Uses
leaf colour lustrous green above, dull green below.
Midrib on adaxial (upper) side narrow (less than 1 No commercially significant uses have been record-
mm), continuous or nearly so, obtusely raised or flat ed of this species. It occurs in remote localities and
and with a central groove, on abaxial side wider, ca. is too small a tree to be of much value for timber,
1 mm, obtusely raised or nearly flat but conspicuous but it may locally have been used for firewood. This
to apex. Stomata small, in numerous intermittent species is not known in cultivation.
lines on either side of abaxial midrib. Pollen cones
axillary, solitary, sessile, elongating to ca. 30 mm,
3 mm wide at anthesis; microsporophylls broadly Podocarpus brassii Pilg., Bot. Jahrb. Syst. 68: 246.
triangular, acute, with two globose pollen sacs. Seed 1937. Pl. 35
cones axillary, solitary on 5–12 mm long, slender pe-
duncles terminating in two minute scales (‘foliola’)
Etymology
subtending the eventually 7–8 mm long, 4–5 mm
wide, succulent red receptacle consisting of an axis This species commemorates the plant collector
with two unequal bracts. Seeds solitary, including L. J. Brass, who made six collecting expeditions to
the epimatium 7–8 mm long, subglobose or broadly New Guinea sponsored by the American Museum
ovoid, with a weakly developed crest. Seed proper of Natural History in New York.
more or less ovoid, 4–5 mm long, grooved or pitted
(in sicco).
Vernacular names
In Flora Malesiana ser. 1, 10 (3): 413–414 (1988) a
Distribution
number of local names for var. brassii are listed; still
Brazil: Bahia, Distrito Federal, Goiás, Mato Grosso, other names were seen mentioned on herbarium
Minas Gerais, Roraima; Venezuela. collections.
 casionally in alpine grassland. It is a high altitude
Description
species; the two varieties, as noted by De Lauben-
Decumbent shrubs or small, often gnarled trees fels in Flora Malesiana ser. 1, 10 (3): 413–414 (1988),
0.15–15 m tall, rarely an erect forest tree to 30 m tall although for the most part sympatric, occur at more
and 75 cm d.b.h. Bark thin and smooth, becom- or less separated altitudes. Var. humilis, a decumbent
ing stringy on largest stems, light brown weather- shrub or small, stunted tree, is found from 2510 m
ing grey; inner bark pink, sometimes nearly white. to 3600 m a.s.l.; the more abundant tree form, var.
Crown of trees often flat-topped, otherwise usually brassii, from occasionally around 2700 m, but more
irregular. Foliage branches spreading to assurgent, commonly between 3100 m and 4000 m a.s.l. In wet
profusely leaved distally, forming dense tufts of foli- areas near pools var. humilis becomes decumbent
age. Terminal buds on leading shoots 3–5 mm wide, and forms dense ‘mats’ of ca. 3 m2 and only 15–30 cm 815
onion-shaped or truncate with triangular, spreading tall. The (fuzzy) forest line is situated at 3600–3700
outer scales. Leaves on young plants 2.5–4 cm long, m in many places, so the highest growing trees of
5–7.5 mm wide, linear-lanceolate, straight, acute or var. brassii are isolated individuals emergent above
short apiculate. Leaves on adult plants (0.5–)1–2(– shrubbery or in acid, boggy subalpine to alpine
2.5) cm long, (3–)4–6(–7) mm wide, oval or elliptical grassland. The species is rather rare in tall forest,
to oblong, coriaceous, spreading to erect and dense- where it may attain 25–30 m with a straight bole. As-
ly crowded, abruptly or in longer leaves gradually ta- sociated taxa in the subalpine shrubbery or mossy
pering to a sessile or short petiolate base; apex obtuse forest are e.g. Rhododendron and Rapanea; in the
or obtuse-cuspidate; margins revolute; midrib raised upper montane forest we can also find the conifers
on adaxial side, forming a narrow ridge less than 0.3 Araucaria cunninghamii, Dacrycarpus compactus
mm wide, ca. 0.5 mm wide and obtusely raised on and Podocarpus pilgeri. Tree ferns are often abun-
abaxial (lower) side; leaf colour in flushing leaves dant in the alpine grassland, which often forms mo-
pinkish or light green with reddish margins, in adult zaic vegetation patterns with shrubbery and patches
leaves dull dark green above, light or glaucous green of mossy forest. The occurrence of P. brassii in grass-
below. Stomata very small, in numerous irregular land away from woody vegetation may be caused by
lines on either side of abaxial midrib. Pollen cones past fires, which are often lighted by people.
axillary, solitary or rarely in pairs, sessile, elongating
at anthesis to ca. 40 mm, slender or robust, 3–8 mm
Uses
wide; microsporophylls short triangular or with an
elongated 2–3 mm long reddish or pink apex, bear- This species is not recorded as being used commer-
ing two globose, greenish yellow pollen sacs. Seed cially; its rarity as a tall tree makes exploitation for
cones towards ends of foliage branchlets, axillary on timber unlikely. Outside a few plants grown in bo-
short peduncles; receptacles subtended by two 3–4 tanic gardens, where it is usually a shrub, but only
mm long bracts (‘foliola’), 8–12 mm long, expanding named to species, neither variety is in cultivation.
at maturity to 15 mm long and wide and becoming Due to their high altitude provenance both forms
dark purple to nearly black when ripe. Seeds includ- should be perfectly hardy for gardens in cool tem-
ing the covering epimatium ovoid, 10–13 × 8–9 mm, perate regions, and the decumbent form of var. hu­
without a crest, olive green or purplish green or pru- milis would make an attractive garden plant worth
inose. Seed proper not observed. being tried.
2 varieties are recognized:
Distribution
Podocarpus brassii Pilg. var. brassii. Type: New
New Guinea.
Guinea: Papua New Guinea, Owen Stanley Range,
TDWG codes: 43 NWG-IJ NWG-PN
Mt. Albert Edward, L. J. Brass 4395 (lectotype K,
here designated). Fig. 256
Ecology
Description
Podocarpus brassii occurs in high montane ever-
green forest, in subalpine mossy forest, in subalpine Small trees, often stunted, to 15 m tall; rarely a tall
to alpine scrubland and on the margins of or oc- tree to 30 m and 75 cm d.b.h. Pollen cones robust,
816 4

2 7

Pl ate 35. Podocarpus brassii. 1. Branch with foliage and pollen cones (var. humilis). 2. Branchlet with
leaves and pollen cones (var. brassii). 3. Branch with foliage and seed cones (var. brassii). 4. Branchlet with
leaves and seed cone (var. brassii). 5. Seed cone. 6, 7. Leaves.
6–8 mm diam. at anthesis; microsporophylls with grey. Crown in more or less sheltered trees conical,
elongated, reddish or pink apex. becoming rounded with age, otherwise usually ir-
regular. Foliage branches spreading to assurgent,
densely leaved distally, forming tufts of foliage;
Distribution
shaded branches with more widely spaced leaves.
New Guinea. Terminal buds on leading shoots 4–6 × 3–5 mm,
TDWG codes: 43 NWG-IJ NWG-PN onion-shaped or truncate with triangular, spreading
outer scales. Leaves on young plants 4–8.5 cm long,
5–9 mm wide, linear-lanceolate, straight, narrowly
Conservation
acute. Leaves on adult plants (1.2–)2–5(–7) cm long,
IUCN: LC 4–6(–8) mm wide, elliptical or falcate-lanceolate to 817
linear-lanceolate, coriaceous, spreading to erect and
Podocarpus brassii Pilg. var. humilis de Laub., Blu- densely crowded, gradually tapering to a sessile or
mea 30 (2): 274. 1985. Type: New Guinea: Papua New short petiolate base; apex acute or obtuse; margins
Guinea, Telefomin District, Mt. Capella, C. Barker slightly revolute; midrib acutely raised on adaxial
LAE 67278 (holotype L). side, forming a narrow ridge less than 0.4 mm wide,
ca. 1 mm wide and obtusely raised on abaxial (lower)
side. Stomata very small, in numerous irregular lines
Description
on either side of the abaxial midrib. Pollen cones ax-
Decumbent shrubs or stunted trees to 5–6 m tall. illary, solitary or rarely in pairs, sessile, elongating at
Pollen cones slender, 3–4 mm wide; microsporo- anthesis to ca. 20 mm, 4–5 mm wide; microsporo-
phylls small, with a triangular apex. phylls with narrowed 1–1.5 mm long apex, bearing
two globose pollen sacs. Seed cones towards ends
of foliage branchlets, axillary on short peduncles;
Distribution
receptacles subtended by two 3–4 mm long bracts
New Guinea. (‘foliola’), 6–8 mm long, distally bilobed or truncate
TDWG codes: 43 NWG-IJ NWG-PN when swollen and becoming dark purple. Seeds in-
cluding the covering epimatium ovoid, 8–10(–12) ×
6–7 mm, with a small distal crest, dark purple, glau-
Conservation
cous or pruinose. Seed proper not observed.
IUCN: LC
Distribution
Podocarpus brevifolius (Stapf) Foxw., Philipp. Malaysia: Sabah (Mt. Kinabalu and surrounding
J. Sci. 6: 160. 1911. Podocarpus neriifolius D. Don mountain ridges).
var. brevifolius Stapf, Trans. Linn. Soc. London, TDWG codes: 42 BOR-SB
Bot., ser. 2, 4: 249. 1894. Type: Malaysia: Sabah,
Ranau District, Mt. Kinabalu, G. D. Haviland 1093
Ecology
(lectotype K, here designated). Fig. 257, 258
Podocarpus brevifolius is locally common in upper
montane to subalpine dwarf forest on Mt. Kinabalu
Etymology
and other mountain ridges and heights in the vicin-
The species epithet refers to the short leaves. ity. It grows most commonly on ultramafic substrate
and also on granite, among boulders or from crevic-
Vernacular names es. It has been found on ultramafic rock at 1350–1450
m a.s.l. near a copper mine and on the Bambangan
No vernacular names are known for this species.
Ridge at 1900 m in lower montane forest. Its altitu-
dinal range on Mt. Kinabalu proper is between 2100
Description
and 3800 m a.s.l., where it is a constituent of a dwarfed
Small, often gnarled trees 1.5–7.5 m tall. Bark be- forest to 10 m tall, dominated by the conifers Phyllo­
coming scaly on largest stems, brown weathering cladus hypophyllus and ­Dacrycarpus ­kinabaluensis
 and the umbrella-crowned angiosperm tree Lepto­ age branchlets terete, finely grooved between leaf
spermum recurvum (Myrtacea). Other common bases, terminating in globose buds with imbricate,
woody plants in this zone are the conifer Dacry­ rounded, keeled scales and apiculate, partly free out-
dium gibbsiae (on ultramafics) and the angiosperms er scales. Leaves on saplings larger than on mature
Eugenia (Syzygium) kinabaluensis, Rhododendron trees, to 5 cm long and 9 mm wide; leaves on mature
buxifolium and Schima brevifolia. Near the summit trees crowded, directed forward at a narrower than
of the mountain only dwarfed, shrub-like plants of 45° angle towards the distal parts of branchlets but
P. brevifolius occur in granite crevices. more widely spreading lower down, 1.5–4 cm long,
straight, elliptic to linear-oblanceolate, (4.5–)6–9
mm wide, gradually tapering to a near sessile to
Conservation
818 short petiolate base, abruptly narrowing to a spines-
IUCN: NT cent apex, the spine 1–1.5 mm long; margins slightly
revolute, texture coriaceous, stiff; leaf colour dark
green above, paler green below. Midrib inconspicu-
Uses
ous on adaxial (upper) side, slightly raised above
No uses are recorded of this species. It can be as- leaf base but soon becoming a groove towards apex,
sumed that provenances from high altitude (above continuous, obtusely raised and wider (0.7 mm) on
3000 m) may be hardy as they experience occasion- the abaxial side. Stomata very small, in numerous
al frost. It will probably be a slow growing shrub in intermittent lines on either side of abaxial midrib.
cultivation, but should be tried. Pollen cones not observed. Seed cones axillary, soli-
tary on 6–8 mm long peduncles; receptacles an axis
with two fused, unequal bracts, growing to 8–10 mm
Podocarpus buchii Urb., Fedde’s Repert. Sp. Nov. long, becoming swollen and succulent, bright red.
Regni Veg. 19 (16–21): 298. 1924. Podocarpus Seeds solitary, elliptical, including the epimatium
angusti­folius Griseb. subsp. buchii (Urb.) Staszk., 7–8 mm long, 4–4.5 mm wide, with a prominent, re-
Fragm. Fl. Geobot. 33: 77. 1988; Podocarpus aris­ curved distal crest. Seed proper not observed.
tulatus Parl. var. buchii (Urb.) Silba, J. Int. Conifer
Preserv. Soc. 7 (1): 31. 2000. Type: Hispaniola: Haiti,
Taxonomic notes
near La Belle­fontaine, L. von Buch 2089b (holotype
B, destroyed?). The leaves of P. buchii var. latifolius in the type speci-
men at K (E. L. Ekman 4913, isotype K) from the
Podocarpus buchii Urb. var. latifolius Florin, Ark. Massif du Nord in Haiti are 1.5–4 cm long and 5–9
Bot. 25-A (5): 3. 1934; Podocarpus angustifolius mm wide. This is overlapping the range of dimen-
Griseb. subsp. buchii (Urb.) Staszk. var. latifolius sions found in specimens of P. buchii Florin ascribed
(Florin) Staszk., Fragm. Fl. Geobot. 33: 77. 1988. to the species, but reaching 2–3 mm wider. This does
not seem to warrant taxonomic distinction, as the
other charcters observed in the type specimen are
Etymology
similar to the species. Gray & Buchholz (1948) in
This species was named after the German botanist their revision of American species of Podocarpus in-
Christian Leopold von Buch (1774–1853). cluded material with much larger leaves in this vari-
ety, which appears to belong to P. aristulatus.
Vernacular names
Distribution
No common names have been recorded for this spe-
cies. Caribbean: Hispaniola: Dominican Republic, Haiti.
TDWG codes: 81 DOM HAI-HA
Description
Ecology
Small trees 5–10 m tall, or larger (?); trunk d.b.h.
to 30 cm (or to 100 cm?). Bark smooth, brown. Podocarpus buchii occurs in the mountain forests
Branches spreading, forming a dense crown. Foli- of Hispaniola. The northern population in Haiti,
­ escribed by Florin (op. cit.) as P. buchii var. latifolius,
d Foliage branchlets terete, with grooves and fine ridg-
occurs on limestone massifs. In most populations in es between the leaf bases, terminating in small buds
the south of this country the geological substrate is with elongated, incurved outer scales. Leaves crowd-
also a hard karst limestone with a lateritic thin soil. ed towards ends of branchlets, spreading forward at
In the Dominican Republic the species is known ca. 35–45° or some lower leaves spreading wider, lin-
from two locations in montane forest on metamor- ear, (1.5–)2.5–5(–7.5) cm long, 2–5 mm wide, straight
phic rock types. The forest is mostly evergreen tropi- or slightly falcate, gradually narrowing to a sessile
cal rainforest dominated by angiosperms with small or short petiolate base, more abruptly converging
leaves on steep, rocky slopes, ridges and summits at distally to an obtuse or minutely acuminate apex;
altitudes between 800 m and 2050 m a.s.l. margins mostly parallel, slightly revolute; texture
coriaceous; leaf colour dark green above, flushing 819
leaves bright glaucous blue. Midrib thin, raised prox-
Conservation
imally but fading towards apex on adaxial (upper)
This species is apparently scattered and can be lo- side, continuous and markedly visible on abaxial
cally common on steep karst formations and moun- side. Stomata small, in numerous intermittent lines
tain ridges. on either side of abaxial midrib. Pollen cones axillary,
IUCN: NT solitary or with 2–3 together on 4–8 mm long pedun-
cles, subtended by scarious bud scales, cylindrical,
elongating to 1–2(–3) cm, 3–4 mm wide at anthesis;
Uses
microsporophylls 1–1.5 mm long and ca. 1 mm wide,
No uses have been recorded of this species. If re- with minutely erose-denticulate margins, each bear-
ports about large trees with d.b.h. to 1 m are correct, ing two subglobose pollen sacs. Seed cones axillary,
such trees may well have been logged and the wood solitary, on a short peduncle, receptacle formed of an
used for general construction, carpentry and furni- axis with two unequal bracts, swelling only slightly
ture making. to 3–4 mm long and 2.5 mm wide. Seeds including
the epimatium 15–18 mm long, 10–12 mm wide, ob-
liquely ovoid with a distal crest, brown to bluish pru-
Podocarpus capuronii de Laub., Adansonia 11 (4): inose or whitish pruinose. Seed proper not observed.
713. 1971. Type: Madagascar: Fianarantsoa Prov.,
Itremo Massif, Mt. Ambatomenaloha, R. Capuron
Taxonomic notes
11566 (holotype P).
Podocarpus woltzii was described by Gaussen (op.
Podocarpus woltzii Gaussen, Bull. Soc. Hist. Nat. cit.) as a distinct species, but with characters said
Toulouse 110: 123. 1974; Podocarpus capuronii de to be close to P. rostratus. The main difference stat-
Laub. var. woltzii (Gaussen) Silba, J. Int. Conifer Pre- ed seems to be the occurrence inside the leaves of
serv. Soc. 7 (1): 31. 2000. horizontal sclereids, hard bodies that are sometimes
interspersed between the mesophyll cells. Sclereids
in leaves of conifers occur erratically and are not
Etymology
reliable taxonomic characters, although some spe-
This species has been named after R. Capuron, who cies may never develop them. Podocarpus woltzii
collected the type specimen. was also described with larger leaves than those of
P. rostratus. The longer and wider leaves seem to be
based on material belonging to P. capuronii de Laub.
Vernacular names
described two years earlier and, like P. woltzii, based
No common names have been recorded for this spe- on a collection made by R. Capuron on the same day
cies. but on a different mountain.

Description Distribution
Shrubs or small trees to 20 m tall, usually less than Madagascar: Itremo Massif, Manandona.
6 m. Bark breaking in small irregular flakes, brown. TDWG codes: 29 MDG
 ing grey. Branches sparse, forming an open, spread-
Ecology
ing crown. Foliage branchlets slender, terete, finely
Podocarpus capuronii is found on skeletal sandy soil grooved between remote leaf bases. Terminal veg-
over quarzite or gneiss, along streams in ravines and etative buds more or less conical, with short, trian-
on rocky slopes and ridges. It is a slow growing shrub gular, acuminate scales 4–6 mm long, 3 mm wide at
or stunted tree in these depauperate sites; in forest base; outer scales free at apex; inner scales more or
it may attain 20 m. The altitude (from information less imbricate. Leaves on saplings and juvenile trees
with herbarium specimens) ranges from 1320–2000 much larger than on mature trees, 12–18 cm long,
m, but is reported to extend to 2800 m a.s.l. 17–23 mm wide, lanceolate-acuminate, thin and
lax, light green. Leaves on mature trees 4.5–7.5 cm
820 long, 8–15 mm wide, elliptic to lanceolate, straight
Conservation
or slightly falcate, more or less abruptly narrow-
Probably now restricted to the Itremo massif, where ing to a short petiolate base, abruptly or gradually
all recent collections have been made. The disjunct tapering to an acute apex; margins flat, mostly not
occurrence is based on Perrier de la Bathie 13163 parallel; texture thin coriaceous; leaf colour deep
in the Paris Herbarium (P), which was collected green above, pale green below. Midrib on adaxial
in the 1920’s (exact date unknown). Habitat loss is (upper) side a shallow groove, fading towards apex;
considered to be very high in the whole of central obtusely raised and continuous to apex on abaxial
Madagascar. Deforestation, habitat loss, cutting for side, ca. 1 mm wide. Stomata very small, in numer-
firewood, fires, and grazing are the main threats to ous intermittent lines forming two broad bands on
this species. This species is not known to occur in a either side of abaxial midrib. Pollen cones axillary,
protected area. solitary, sessile, subtended by small, triangular peru-
IUCN: EN (A2c, A3c) lar scales, cylindrical, 20–35 mm long and creamish
white when mature. Seed cones axillary, solitary on
a 4–10 mm long peduncle; receptacles 7–9 mm long,
Uses
slightly swollen and red when mature. Seeds includ-
No uses have been recorded of this species. It is likely ing the epimatium subglobose to globose, 9–12 mm
to have been used as firewood when still abundant; long, 8–10 mm wide, with a small apiculus, green.
perhaps the wood of larger trees was used in local Seed proper globose, with a lustrous, finely pitted,
construction and hand tool carpentry. brown seed coat.

Taxonomic notes
Podocarpus celatus de Laub., Fl. Venezuela 11 (2):
35. 1982. Type: Bolivia: Potosí, Morro, R. W. Pearce David de Laubenfels (op. cit.) cited as the holotype
s.n. (lectotype K, specimens B, designated here). of Podocarpus celatus “Pearce en 1866 (K), Bolivia,
Moro.” The pertinent sheet at Kew contains a foli-
age branch with large leaves from a juvenile tree
Etymology
(Pearce: “leaves of young trees very large”) and two
The species epithet (Latin: celatus = concealed, hid- branchlets, one fertile with a seed cone, with much
den) relates to the species previously not being rec- smaller leaves from a mature tree. These are in ef-
ognized as distinct from P. magnifolius. fect two separate gatherings and cannot be both the
holotype. I have marked them A (juvenile plant) and
B (mature plant) and here designate the two branch-
Vernacular names
lets marked B as the lectotype of this species. The
’Cinqui-masé’ (Peru) juvenile foliage branch thereby becomes a paratype.
This species differs in its foliage only slightly from
P. magnifolius, with which the type specimens at Kew
Description
of P. celatus were provisionally equated by J. T. Buch-
Trees to 25–30 m tall; trunk to 60 cm d.b.h., erect holz in 1950. Several leaves of the juvenile foliage
in tall forest habitat. Bark thin, fibrous, exfoliating specimen on the ‘type sheet’ at K have acuminate
in small strips and flakes, reddish brown, weather- leaves, similar to leaves of P. magnifolius. The buds of
P. celatus have shorter scales and the seeds are nearly
Description
globular without a crest. The pollen cones, unknown
when P. celatus was published, are similar to those of Small trees to 8 m tall; trunk to 20 cm d.b.h. Bark
P. magnifolius, but slightly shorter at ­maturity. smooth, fibrous, brown weathering to grey-brown.
Branches ascending, foliage branches in mature
plants dense, forming a (broad) columnar crown; ul-
Distribution
timate branchlets angular, glabrous, terminating in a
Bolivia: Potosí; Brazil: Mato Grosso; Colombia; small conical bud with carinate, somewhat tapered,
Peru: Amazonas, Junin, Loreto, Montaña, Puno; to 3 mm long and occasionally spreading scales.
Venezuela: Amazonas, Bolivar, Tachira. Leaves densely crowded, spirally arranged, spread-
TDWG codes: 82 VEN 83 BOL COL PER 84 BZC-MT ing to erect and rigid, sessile to short petiolate, leaf 821
blade linear-lanceolate to oblanceolate, 1.2–3.5 cm
long, 2.5–4.5 mm wide, gradually tapering to base,
Ecology
in adult leaves with an obtuse apex, midvein acutely
Podocarpus celatus occurs in tropical evergreen low- raised adaxially, not raised and wider abaxially; leaf
land to low montane rainforests. Its altitudinal range colour olive green above (adaxially), glaucous green
as presently known from herbarium specimen data below. Stomata conspicuous, in numerous regular
is between 350 m and 1400 m a.s.l. lines on abaxial side on either side of the midvein.
Pollen cones axillary, sessile, in clusters of (1–)2–3
per axil, subtended by several scarious, more or less
Conservation
triangular bracts, long cylindrical, 15–25 mm long,
IUCN: LC 1.5–2 mm wide; microsporophylls spirally arranged,
imbricate, spreading at anthesis, the laminar part
apiculate, 0.5–0.7 mm long, with two basal pollen
Uses
sacs. Seed cones and seeds not known.
This species is undoubtedly being logged for its tim-
ber, as it grows in accessible forests to a considerable
Taxonomic notes
size. As with other wood of podocarps, it is used for
general construction, carpentry and furniture mak- This species was originally described as a variety of
ing. It is not known to be in cultivation, or perhaps the wide-spread species Podocarpus macrophyllus
planted in a few botanic gardens as P. magnifolius or (Thunb.) Sweet by N. E. Gray (op. cit.), based on the
P. oleifolius. collection R. C. Ching 2477 of which she had seen
a specimen in the Herbarium of the Arnold Arbo-
retum at Cambridge, Massachusetts. S. Y. Hu (op.
Podocarpus chingianus (N. E. Gray) S. Y. Hu, cit.) described it as a new species, based on the same
Taiwania 10: 32. 1964. Podocarpus macrophyllus type collection (R. C. Ching 2477) so in fact elevating
(Thunb.) Sweet var. chingii N. E. Gray, J. Arnold Gray’s variety in rank to a species. Two more collec-
Arbor. 39: 474. 1958; Margbensonia chingiana tions were cited by Hu (Gray only knew the type),
(N. E. Gray) A. V. Bobrov & Melikyan, Bjull. Mosk­ C. L. Tso 1477 from Jiangsu and W. P. Fang 14261
ovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998. from Sichuan. A duplicate of C. L. Tso 1477 is in the
Type: China: Zhejiang, [“55 Chinese miles (li) west of Kew Herbarium (K); the collection from Sichuan
Lung-sien”], R. C Ching. 2477 (holotype A). Fig. 259 could not be verified. This taxon was regarded as a
cultivar of Podocarpus macrophyllus (cv. Chingii) by
S. Y. Zhang in Flora of Zhejiang Province 1 (Lin &
Etymology
Zhang, 1993), but there is no evidence of it having
The specific and varietal epithets commemorate the been found (only) in cultivation.
Chinese plant collector Ching Ren-chang.
Distribution
Vernacular names
China: Jiangsu, Zhejiang, Sichuan(?).
zhu guan luo han song (Chinese) TDWG codes: 36 CHS-JS CHS-ZJ
 becoming rounded with age in large trees, other-
Ecology
wise usually irregular or stunted. Foliage branches
Podocarpus chingianus occurs in woods and open spreading, mostly with widely spaced leaves. Termi-
thickets, i.e. secondary woodland or shrubby vegeta- nal buds on leading shoots 6–10 mm long, 3–4 mm
tion. Its altitudinal range is quoted as “near sea level wide, narrowly conical, with erect, free, long acumi-
to 1000 m”, but this is based on no more than two nate outer scales. Leaves on young plants 10–15(–20)
(or three) localities cited in the literature. This habi- cm long, 9–12 mm wide, linear-lanceolate, straight
tat note is quoted from S. Y. Hu (1964), but was not or slightly curved, acute. Leaves on adult plants
specified with the collected specimens from Jiangsu smaller (3.5–)5–10(–12) cm long, (5–)7–10(–12) mm
and Zhejiang (the Sichuan specimen was not seen). wide, lanceolate to linear-lanceolate, coriaceous,
822 It is apparently very rare. spreading to ascending, gradually tapering to a peti-
olate base; apex acute, or slightly apiculate from ex-
tended midrib; margins flat; midrib obtusely raised
Conservation
and 0.8–1 mm wide on adaxial side, flattened and
Too little is known about this taxon, apart from its slightly wider on abaxial (lower) side. Stomata very
apparent rarity, to evaluate its conservation status. It small and inconspicuous in dense lines on abaxial
is with certainty known from two localities, one in side. Pollen cones axillary, solitary or occasionally
Jiangsu and the other in Zhejiang; a third herbarium in pairs, sessile, subtended by long acuminate bracts
collection was mentioned by S. Y. Hu (op. cit.) to be (bud scales), elongating at anthesis to 20–40 mm,
from Sichuan. That would be far to the west of the 3–4 mm wide; microsporophylls with acute apex ca.
other two, neither of which has been specified to 1 mm long, bearing two globose pollen sacs. Seed
modern topographic locality names, so they remain cones axillary on a 4–13 mm long peduncle; recep-
essentially located as to the provinces only. tacles subtended by a 3–5 mm long bract, 8–12 mm
IUCN: DD long, formed of an axis with two unequal parts, swell-
ing to become succulent and red. Seeds including the
epimatium ovoid-globose, ca. 10 × 7 mm when ma-
Uses
ture, distally obtuse. Seed proper not observed.
No uses are known of this species; it may be in culti-
vation in China locally.
Distribution
Malaysia: Sabah (Bukit Ampuan, Mt. Silam),
Podocarpus confertus de Laub., Blumea 30 (2): Sarawak.
271. 1985. Type: Malaysia: Sabah, Tawau District, TDWG codes: 42 BOR-SB BOR-SR
Gunung Silam, D. J. de Laubenfels P 691 (holotype L).
Ecology
Etymology
Podocarpus confertus occurs in kerangas forest on
The species epithet means ‘pressed close together’, poor, sandy soils and in stunted forest on ultrabasic
but what is considered to be so is not clear from the rocks, commonly at altitudes between 100 and 1200
original description or the specimens seen. m a.s.l. It can locally form dense populations, but
commonly it is associated with various shrubs and
trees, conifers as well as angiosperms.
Vernacular names
No common names have been recorded for this spe-
Conservation
cies.
IUCN: LC
Description
Uses
Trees to 35 m tall; trunk erect, terete, to 70 cm d.b.h.
Bark soft, becoming flaky on largest stems, peeling in Where it grows to a large tree this species will be
longitudinal strips, brown weathering grey. Crown valued and exploited for its timber; however, most
trees do not attain large size as they occur on very of leaf, but fading towards apex, often grooved (in
poor soils or on exposed sites with ultrabasic rock. sicco), on abaxial side continuous to apex, wider (ca.
Its wood may be used for house construction, floor- 1 mm), but flattened. Stomata small, in numerous
ing, furniture making and cabinet work, carpentry intermittent lines on either side of abaxial midrib.
and joinery, household utensils, matches, and tooth- Pollen cones axillary, solitary or sometimes in pairs,
picks. This species is not known to be in cultivation. sessile, subtended by scarious bud scales, long cy-
lindrical, elongating to 50–60 mm, 3–4 mm wide at
anthesis; microsporophylls ovate-triangular, obtuse
Podocarpus coriaceus Rich., in A. Richard (ed.) with erose margins, each bearing two globose pollen
Comm. Bot. Conif. Cycad.: 14, t. 1, f. 3. 1826. Type: sacs. Seed cones axillary, solitary on 4–8 mm long
Caribbean: Montserrat. R. Brown s.n. (holotype peduncles, 2 bracts at base of receptacle (‘foliola’) 823
BM). soon absent; receptacles composed of an axis with
two unequal, fused bracts with free tips, swelling to
a 7–9 mm long, 6–7 mm wide, succulent red body.
Etymology
Seeds solitary, including the epimatium 8–10 mm
The species epithet (Latin corium = skin) describes long, 6–7 mm wide, obliquely ovoid with a sub­apical
the thick, leathery leaves. beaked crest, green. Seed proper not observed.

Vernacular names Distribution

Resinier moutaigue (Dominica); Caoba del país, Caribbean: Dominica, Guadeloupe, Martinique,
Coabilla (Puerto Rico); Granadillo (Trinidad & To- Montserrat, Puerto Rico, Saint Lucia, Saint Kitts and
bago) Nevis, Trinidad and Tobago.
TDWG codes: 81 LEE-GU LEE-MO LEE-SK PUE TRT
WIN-DO WIN-MA WIN-SL
Description
Shrubs or more commonly small, stunted trees
Ecology
3–10 m tall, rarely to 20 m; d.b.h. to 50 cm. Bark
thick, smooth, becoming fissured and scaly, exfo- Podocarpus coriaceus occurs in lowland forest or
liating with shaggy strips, brown weathering grey. woodland on poor sandy soils in Trinidad and To-
Branches much spreading and contorted in older bago and on the Leeward and Windward Islands of
trees. Foliage branchlets stout, terete, with longitu- the Lesser Antilles and in Puerto Rico in ‘elfin forest’
dinal grooves and ridges on vigorous shoots, less on windswept mountain ridges and summits from
marked on slow growing shoots, terminating in 500 m to 1100 m a.s.l. This vegetation type does not
large, broadly ovoid buds with ovate-apiculate 3–6 exceed 10 m in height and is usually much lower and
mm long scales which are free towards apex, a few scrubby.
of the outer scales sometimes elongated to scale-
like leaves 10 mm long or longer. Leaves on vigor-
Conservation
ous shoots of saplings larger than on mature trees,
to 22 cm long and 17 mm wide. Leaves on mature IUCN: LC
trees (3–)5–12 cm long, 7–14 mm wide, straight or
slightly falcate, linear-lanceolate to linear, narrow-
Uses
ing to a near sessile or short petiolate base, gradually
tapering to an acute or slightly acuminate apex, only No uses have been recorded of this species; where
in longer leaves the two sides running parallel in the accessible it is probably cut for firewood. In the past,
middle section of the leaf; margins mostly straight, larger trees to 18 m tall may have existed, which were
sometimes slightly revolute; texture coriaceous; leaf logged long ago and were probably used for con-
colour dark green above, dull green below. Midrib struction and general carpentry.
on adaxial (upper) side thin, raised in proximal part
 Podocarpus costalis C. Presl, Epimel. Bot.: 236. 6–7 mm, with a small distal crest, green or glaucous.
1851. Type: [orig. mat. coll. T. Haenke (1792), not Seed proper not observed.
located]. Fig. 260
Distribution
Podocarpus costalis C. Presl var. taiwanensis Gaus-
sen, Trav. Lab. Forest. Toulouse T. 2, 1 (2, 21): 183. Philippines: N Luzon coast?, Batan to Bucas Islands
1976 (nom. inval., Art. 37.1). in the Luzon Strait; Taiwan: Lanyu (Orchid Island).
TDWG codes: 38 TAI 42 PHI
Etymology
Ecology
824 The species epithet refers to the coastal habitat of
this species. Podocarpus costalis occurs on islands in lowland ev-
ergreen scrub to low forest on karst limestone bluffs
and ridges or sea-stacks, from near sea level to at
Vernacular names
least 300 m a.s.l. It may be native in similar habitat
lan yu luo han song (Chinese); no common names on the northern coast of Luzon.
recorded in the Philippines.
Conservation
Description
Podocarpus costalis has a very limited distribution in
Small, often gnarled trees 1.5–10 m tall. Bark becom- fewer than five localities (islands) and is restricted
ing scaly on largest stems, brown weathering grey. to a specific habitat. It is in decline because plants
Crown usually spreading, dense and irregular. Foli- are taken from the wild to be planted in gardens in
age branches spreading to assurgent, densely leaved Luzon and Taiwan, and possibly elsewhere. It was
distally, forming tufts of foliage; shaded branches listed on Appendix I of CITES in 1975, but deleted
with more widely spaced leaves. Terminal buds on from the Appendices in 1990. Attempts should be
leading shoots 3–4 mm wide and 2–4 mm long, made to grow this species from seed or scions in or-
onion-shaped or truncate with triangular, erect der to minimize the taking of plants from habitat.
and free outer scales. Leaves on young plants to 9 IUCN: EN (A1c)
cm long, 10–13 mm wide, linear-lanceolate, straight,
acute or more or less rounded at apex. Leaves on
Uses
mature plants (2–)3–5(–7) cm long, 5–8(–10) mm
wide, oblong to linear, coriaceous, spreading to erect This attractive, shrubby dwarf tree is popular in cul-
and densely crowded, gradually tapering to a short tivation for gardens in the Philippines and in Taiwan,
petiolate and twisted base; apex obtuse to rounded, where it has been confused with P. polystachyus. The
sometimes indistinctly apiculate; margins slightly pollen cones of P. costalis are much more robust (up
revolute; midrib acutely raised on adaxial side, to 3 times wider) and its leaves shorter and more
forming a narrow ridge less than 0.4 mm wide, ca. crowded. The ripe receptacle of the seed cone is dark
1 mm wide and obtusely raised on abaxial (lower) purple-pruinose, not red. Podocarpus costalis is also
side. Stomata very small, in numerous irregular used in bonsai culture.
lines on either side of abaxial midrib. Pollen cones
axillary, solitary, sessile with rounded bud scales at
base, (short) cylindrical, 8–25 mm long and 6–8 mm Podocarpus costaricensis de Laub. ex Silba,
wide at anthesis; microsporophylls with narrowed Phytologia 68: 67. 1990. Type: Costa Rica: San José,
ca. 2 mm long apex, bearing two globose pollen sacs. Tarrazu, San Pablo de Leon Cortes, 2 km W of
Seed cones towards ends of foliage branchlets, axil- cemetery, D. J. de Laubenfels P 810 (holotype MO).
lary on slender peduncles; receptacles subtended by
two 1–1.5 mm long, deciduous bracts (‘foliola’), 10–14
Etymology
mm long, distally bilobed or truncate when swollen
and becoming reddish purple then dark purple-pru- The species epithet indicates Costa Rica, where the
inose. Seeds including the epimatium ovoid, 8–10 × species is endemic.
 is disjunct from P. matudae, which has not been re-
Vernacular names
corded with certainty south of Guatemala, and at
No common names have been recorded for this spe- present remains only known from a limited area in
cies. Costa Rica; De Laubenfels mentions an additional
locality (Parque Nacional de Darién) in Panama, but
no specimens.
Description
Trees to 20–30 m tall, erect, monopodial, with a trunk
Distribution
to 1 m d.b.h. but most known trees are more slender.
Bark reddish brown, scaly and exfoliating in longi- Costa Rica (San José).
tudinal strips; inner bark pinkish. Branches spread- TDWG codes: 80 COS 825
ing, forming a rounded crown. Foliage branchlets
with prominent leaf scars and fine grooves from
Ecology
decurrent leaf bases. Terminal vegetative buds with
spreading acuminate scales 10–15 mm long; axillary Podocarpus costaricensis occurs in lowland and low-
buds small, with triangular, keeled, acute to acumi- er montane tropical rainforest at elevations between
nate scales 2–3 mm long, with free tips. Leaves on ju- 70 m and 1700 m a.s.l. Its distribution and ecology
venile plants larger than on mature plants. Leaves on remain poorly known since it was described in 1990.
mature trees lanceolate, flat, (4–)6–10(–13) cm long,
(7–)10–18(–22) mm wide, straight, narrowing to a
Conservation
petiolate base, gradually tapering to a mostly acu-
minate apex. Midvein lying in a groove on the adax- This species is limited to four localities in Costa
ial (upper) side, narrow (less than 0.5 mm wide), Rica within an extent of occurrence (EOO) of less
extending from base to apex, prominently raised than 10,000 km2. The altitudinal range of 70 m to
and 1–1.5 mm wide on abaxial side and extending 1650 m a.s.l., taken from herbarium collections, in-
to apex. Stomata small, in numerous intermittent dicates a probable decline because of deforestation
lines on either side of midrib on abaxial side only. and changes in land use at lower levels. The locality
Pollen cones axillary, solitary, sessile, subtended by at 70 m was recently collected (1995). Only eight col-
triangular, acuminate, keeled bud scales, cylindri- lections were found in the search of herbaria, indi-
cal, expanding to 20–55 mm long, 4–5 mm wide at cating that the species is uncommon. This species is
anthesis; microsporophylls triangular-acuminate, not recorded from any protected area.
with two basal pollen sacs. Seed cones axillary, not IUCN: VU [B1, a, b (iii)]
observed (the receptacle presumably succulent and
red at maturity).
Uses
No economic uses are known of this species and it is
Taxonomic notes
not in cultivation.
According to De Laubenfels (in Silba, 1990) this
species is the only species in Costa Rica with long
acuminate vegetative bud scales; its leaf apices are Podocarpus crassigemma de Laub., Blumea 26
also (mostly) acuminate. The only other species with (1): 141. 1980 [“crassigemmis”]. Type: Papua New
similar bud scales in Mesoamerica is P. matudae; its Guinea: Eastern Highlands, Goroka, Marafunga,
leaves are usually gradually tapering to an acute apex D. J. de Laubenfels P 743 (holotype L).
and only occasionally acuminate; they are also often
narrower but there is considerable variation in this.
Etymology
The midvein has characteristics similar to that spe-
cies. Although now known from a few more collec- The species epithet (Latin crassus = thick, gemma
tions than the two made by De Laubenfels, the seed = bud) refers to the stout buds on leading foliage
cones of P. costaricensis are still unknown; they are shoots.
probably similar to those of P. matudae. The ­species
 Vernacular names Ecology
Numerous names are known for this species in the Podocarpus crassigemma is a common large canopy
New Guinea Highlands; several are listed in Flora tree in montane rainforest with other Podocarpus
Malesiana, ser. 1, 10 (3): 413 (1988). These names spp. and angiosperms of the families Cunoniaceae,
are likely to be applied to P. archboldii and/or other Fagaceae, Lauraceae, Myrtaceae, and others. It can
podo­carps as well. be an emergent but smaller tree in upper montane
mossy forest with Nothofagus spp., Myrtaceae and
the conifer Phyllocladus hypophyllus, and may ex-
Description
tend into the subalpine acid grassland zone with
826 Trees to 40 m tall, close to tree line only 3–8 m tall; Papuacedrus papuana and tree ferns. Its altitudinal
trunk erect, to 1 m d.b.h., sometimes fluted from range is from (1800–)2100 m to 3400 m a.s.l. and it
base. Bark thin, longitudinally fissured and scaly occurs most commonly on igneous or metamorphic
on largest trunks, fibrous light brown; inner bark rock with acidic soils.
light orange. Crown usually spreading, open and ir-
regular, on exposed ridges sometimes flat-topped.
Conservation
Foliage branches spreading to assurgent, stout; ter-
minal buds on leading shoots (3–)4–8 mm wide and IUCN: LC
4–7 mm long, onion-shaped or truncate with lan-
ceolate, (strongly) recurved outer scales. Leaves on
Uses
young plants to 20 cm long, 10–13 mm wide, linear-­
lanceolate, straight, acute at apex. Leaves on adult Podocarpus crassigemma is a valuable timber tree
plants (2.5)4–9(–11) cm long, 5–8(–10) mm wide, where it attains large sizes with a clear, straight bole.
elliptic to oblanceolate, coriaceous; margins slightly Its wood is used as roundwood for masts, spars and
revolute, gradually tapering to a petiolate base; apex poles, in house construction as beams, in high-
acute or obtuse, sometimes indistinctly apiculate; grade construction for flooring, joinery and other
midrib acutely raised on adaxial side, forming a nar- carpentry, for furniture and cabinet work, veneer, to
row ridge less than 0.4 mm wide, ca. 1 mm wide and make boxes, and for match sticks. Its traditional uses
obtusely raised on abaxial (lower) side; leaf colour include village house construction, household uten-
lustrous dark green above, young leaves flushing yel- sils and wood carving. This species was formerly
low-green, glaucous or red. Stomata very small, in confused with P. archboldii (e.g. Van Royen, Alpine
numerous irregular and intermittent lines on either Flora of New Guinea 2: 30, t. 41. 1979) and is prob-
side of abaxial midrib. Pollen cones axillary, solitary ably not distinguished from it by Papuans, who may
or sometimes in pairs, on a 3–6 mm long peduncle, use either species for the same purposes.
with rounded bud scales at base, (short) cylindrical,
10–25 mm long and 6–7 mm wide at anthesis; mi-
crosporophylls with narrowed ca. 2 mm long apex, Podocarpus cunninghamii Colenso, Visit Ruahine
bearing two globose pollen sacs. Seed cones axillary Mt. Range: 58. 1884. Type: New Zealand: [locality
on slender peduncles; receptacles subtended by two unknown], W. Colenso 1631 (lectotype K, designated
1–1.5 mm long, deciduous bracts (‘foliola’), 13–18 here). Fig. 261
mm long, truncate when swollen and becoming red
then dark purple to shining black. Seeds including Podocarpus hallii Kirk, Forest Fl. New Zealand: 14.
the covering epimatium ovoid, 10–14 × 9–10 mm, 1889; Podocarpus totara G. Benn. ex D. Don var.
with a small distal crest, green or glaucous pruinose. hallii (Kirk) Pilg., in Engler, Pflanzenr. IV.5 [18]: 84.
Seed proper not observed. 1903.

Distribution Etymology
Papuasia: New Guinea (central highlands), Bis- This species was named after Allan Cunningham
marck Archipelago (New Ireland). (1791–1839), who botanized in Australia and New
TDWG codes: 43 BIS NWG-IJ NWG-PN Zealand.
 literature, including earlier handbooks of conifers.
Vernacular names
Brian Molloy in 1991 provisionally selected a sin-
Hall’s totara, montane totara gle branchlet (which he marked with ‘a’) of Colenso
1631 at Kew (K) as the lectotype of P. cunninghamii
Colenso but never came round to publish that
Description
choice. There are three branchlets plus a packet with
Trees to 20 m tall, with a straight trunk to 1.3 m broken material on the sheet at K, all likely to have
d.b.h. Bark thin, scaly, exfoliating in thin, long strips, come from the same tree, and I take the view that
light orange-brown. Branches spreading, forming therefore the entire sheet can serve as the lectotype,
a dense, rounded crown. Foliage branchlets terete, which is here designated as such.
glabrous, finely grooved; branching irregularly, ulti- 827
mate branchlets more or less pectinate, terminating
Distribution
in small, subglobose buds with imbricate, acute or
rounded scales. Leaves helically arranged, sessile or New Zealand: North Island, South Island, Stewart
short petiolate, short decurrent, spreading at wide Island.
angles, lanceolate to lanceolate-linear, (1.5)2–3(–3.5) TDWG codes: 51 NZN NZS
cm long, 3–5(–6.5) mm wide (on saplings and young
trees to 5 cm long), straight or slightly curved, taper-
Ecology
ing to a narrow, twisted base and an acute-pungent
or sometimes obtuse-mucronate apex, coriaceous, Podocarpus cunninghamii occurs in evergreen conif-
stiff; midvein a narrow groove through the middle erous and mixed forest from near sea level to 1000
adaxially (upperside) and an obtuse midrib abaxi- m. In lowland forests up to ca. 500 m a.s.l. P. cunning­
ally; leaf colour mid green, slightly paler green be- hamii may occur together with P. totara; the latter
low. Stomata in two broad bands of intermittent rarely occurs above that elevation. Other common
lines on abaxial side. Pollen cones axillary, solitary and large conifers in these lowland forest remnants
or with 2–5 on distinct peduncles, subtended by are Agathis australis (in Northland), Dacrycarpus
small, triangular scales, cylindrical, 20–30 mm long, dacrydioides, Dacrydium cupressinum, and Prumno­
5–8 mm wide at anthesis, straight or curved after pitys ferruginea, locally Halocarpus kirkii, Manoao
shedding pollen, yellowish green turning brown; colensoi and Phyllocladus trichomanoides. In mon-
micro­sporophylls helically arranged, imbricate, pel- tane situations above 600 m conifers are fewer and
tate-triangular, with denticulate upper margin and angiosperms like Metrosideros umbellata, Quintinia
with two sublateral, globose, yellow pollen sacs. Seed acutifolia and Weinmannia recemosa become more
cones axillary, solitary on a short peduncle; recepta- prominent. Here other conifers are Libocedrus bid­
cles when growing narrowly oblong, with two distal willii, Halocarpus biformis and Phyllocladus tri­
bracts subtending 1–2 ovules, swelling to 4.5–6 mm chomanoides var. alpinus. Especially in South Island
long and 3.5–4 mm wide, yellow ripening to scarlet, Nothofagus solandri often forms dominant patches,
soft and succulent. Seeds including the epimatium but these are not long-lived and Libocedrus bidwillii
ovoid-oblong, with a constricted distal part, 3.5–4 × and Podocarpus cunninghamii are both able to pre-
2–2.5 mm, light green or glaucous green. Seed prop- vail as emergents.
er ca. 3 mm long, narrowly ovoid, seed coat smooth
and hard.
Conservation
IUCN: LC
Taxonomic notes
Podocarpus hallii, a species described by Thomas
Uses
Kirk (op. cit.), has turned out on careful examina-
tion of types and other material to be synonymous The use of Hall’s (or montane) totara as a timber
with the earlier described species P. cunninghamii. tree has virtually vanished with the legal protection
Colenso (op. cit.) published his species in a taxo- of native trees against logging in New Zealand. Al-
nomically obscure title, so it was often overlooked though potentially not as big a tree, its use would
and the name coined by Kirk appears in much of the have been very similar to that of totara (P. totara).
 Podocarpus decumbens N. E. Gray, J. Arnold Arbor.
Taxonomic notes
36: 202. 1955. Type: New Caledonia: Grande Terre,
Province Sud, Montagne des Sources, L. Chevalier The botanical characteristics, apart from growth
s.n., 1949 (holotype ILL). habit, are similar to those of P. longefoliolatus, but
only single-seeded cones (receptacles) are reported
for this decumbent taxon. The number of localities
Etymology
known and herbarium collections made are limited
The species epithet means ‘prostrate, lying on the and it is necessary to expand the field knowledge of
ground’; such plants only raise the growing tip of this peculiar species of Podocarpus.
the shoots.
828
Distribution
Vernacular names
New Caledonia: Grande Terre, southern mountains.
No common names have been recorded for this spe- TDWG codes: 60 NWC
cies.
Ecology
Description
Podocarpus decumbens occurs in open woodland
Decumbent or prostrate shrubs, sometimes semi- or scrubby dwarf forest on mountain ridges at 800–
erect, rooting from stems, spreading over rocks or 1000 m a.s.l., where it forms extensive colonies by
other shrubs. Bark thin, exfoliating with fibrous layering. On open rocky terrain it does not normally
strips; outer bark yellowish brown; inner bark red- rise above 40 cm, but with the support of other veg-
dish brown. Foliage branches terete, finely striated; etation it can erect itself higher above the ground.
terminal buds ca, 4 mm long, with erect lanceolate It is not in any way a liane, as De Laubenfels (1972)
outer scales to 10 mm long, but inner scales much suggested, because it neither entwines other plants,
shorter. Leaves often in tufts towards ends of brach- nor has it developed specialized climbing or holding
lets, or inserted more remotely in shaded shoots, devices or liane architecture (see Bell, 1991).
oblanceolate (widest above the middle part) to line-
ar, 4.5–8 cm long, 6–9 mm wide, coriaceous, gradu-
Conservation
ally tapering to a petiolate base; margins revolute,
straight; apex narrowing abruptly, obtuse or some- Based on the very limited number of localities (only
times acutish. Midrib prominent, acute or obtuse two are known thus far), this species is at risk. Due
on adaxial (upper) side, flattened on abaxial side, to fire hazard the quality of habitat is projected to
both ca. 1 mm wide. Leaf colour light green on both decline, probably leading to a reduction of popula-
surfaces. Stomata small, in numerous intermittent tion size.
lines forming bands on either side of abaxial mid- IUCN: CR [B2ab(iii)]
rib. Pollen cones axillary, solitary, sessile, subtended
by triangular, keeled bud scales, elongating to 15–30
Uses
mm, 3–3.5 mm wide at anthesis; microsporophylls
spirally arranged, triangular, bearing two sub-basal No uses have been recorded of this species. Its inter-
pollen sacs. Seed cones axillary, solitary on a 5–7 esting habit may indicate potential in horticulture in
mm long peduncle; receptacles subtended by two countries with a warm, frost-free and humid climate.
3–4 mm long, linear bracts (‘foliola’), composed of
an axis with 3–4 fused bracts with exserted tips, one
Podocarpus deflexus Ridl., Fl. Malay Pen. 5: 283.
of which is fertile, all swelling to a 8–10 cm long, red
1925. Type: Malaysia: Peninsular Malaysia, Pahang,
and succulent body. Seeds solitary, including the
Gunung Tahan, H. N. Ridley 16024 (lectotype K).
epimatium 6–7 × 4 mm (probably growing slightly
larger but no completely mature seeds are known),
Etymology
with a small crest. Seed proper not observed.
The species epithet refers to the leaves and outer bud
scales being abruptly turned downward (deflected).
 Vernacular names Conservation
No vernacular names are known for this species. This species is known only from two greatly disjunct
populations, the one in Sumatera only known from
one herbarium collection made in 1972. This species
Description
appears to be confined to summit areas of moun-
Small trees to 10 m tall, to 20 cm d.b.h., sometimes tains. Based on these two locations its area of oc-
monoecious. Bark smooth, thin, brown; inner bark cupancy (AOO) is estimated to be less than 500 km2
reddish brown, fibrous. Branches spreading, form- and is probably in the lower range between Endan-
ing a rounded or irregular crown. Foliage branchlets gered (EN) and Critically Endangered (CR) (10–500
with distinct decurrent ridges from the leaf bases, km²). There are no collections known in institution- 829
glabrous, terminating in broad, 2–3 mm long buds al herbaria made after 1973; all were made within
with spreading to reflexed, lanceolate outer scales protected areas, yet habitat degradation remains a
to 10–12 mm long and much shorter, rounded in- problem in these areas, mainly through fires. The
ner scales. Leaves on juvenile plants petiolate, linear, known collections were recorded from the Taman
20–26 cm long, 12–15 mm wide, gradually narrowed Negara (Malaysia) and Gunung Leuser Reserves.
at base and apex, acute. Leaves on mature trees IUCN: EN [B2, a, b (iii)]
nearly as long but narrower, linear, 10–22 cm long,
8–11(–13) mm wide, straight or (irregularly) curved,
Uses
thick coriaceous, deflected at the gradually nar-
rowing, petiolate base so as to all hang downward, No economic uses have been recorded of this rare
tapering to an obtuse or sometimes acute apex. Mid- species.
vein acutely (sometimes obtusely) raised adaxially,
ca. 1 mm wide, flat, marginally wider and drying to
a shallow groove abaxially. Stomata very small, in Podocarpus dispermus C. T. White, Contr.
numerous irregular lines on abaxial (under) side on Arnold Arbor. 4: 10. 1933. Margbensonia disperma
either side of midrib. Pollen cones axillary, in clus- (C. T. White) A. V. Bobrov & Melikyan, Bjull.
ters of 2–3, (nearly) sessile with several basal bract Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1):
scales, cylindrical, elongating to 25–40 mm, 2–3 mm 60. 1998. Type: Australia: Queensland, Atherton
wide; microsporophylls spirally arranged, triangu- Tablelands, Gadgarra Forest Reserve, S. F. Kajewski
lar, spreading, with two basal pollen sacs. Seed cones 1192 (holotype BRI). Fig. 262
axillary, solitary on a 5–10 mm long peduncle; recep-
tacles with 2 subulate bracts (‘foliola’) 2–5 mm long
Etymology
at base and 2 fertile and 1 sterile bracts distally, swell-
ing to become ellipsoid with a truncate distal end, The species epithet refers to the two ovules (seeds)
bright red and succulent when ripe. Seeds enclosed often found on a single receptacle.
in a smooth epimatium, ovoid to subglobose, 10–12
mm long, 6–8 mm wide. Seed proper not ­observed.
Vernacular names
Distribution No common names have been recorded for this spe-
cies.
Malaysia: Gunong Tahan in Pennisular Malaysia;
Indonesia: N Sumatera (Aceh).
TDWG codes: 42 MLY-PM SUM Description
Trees to 20–25 m tall; trunk erect, terete, to 40 cm
Ecology
d.b.h. Bark thin, smooth, on larger trunks becom-
Podocarpus deflexus is a small tree rising above ing flaky, exfoliating in narrow strips, light brown
and locally dominant in dwarf mountain scrub at to grey-brown; inner bark slightly fibrous, pinkish.
altitudes between 1500 m and 2100 m a.s.l. Field Branches spreading, forming a broad crown. Foliage
­observations appear to indicate that this species is branchlets smooth or with fine grooves, terminat-
monoecious, unlike most species in the genus. ing in small, narrowly conical buds with ­lanceolate,
 minutely denticulate, apically free, spreading or Threatened. Stochastic events like cyclones and fires
slightly recurved scales. Leaves on juvenile and are the main threats to this species. Only a limited
mature trees similar in size ranges, spreading at part of the total population occurs within the ‘Wet
a forward angle from branchlets, short petiolate, Tropics of Queensland’ World Heritage Site; most
(5–)8–18(–20) cm long, (13–)18–30 mm wide, the known trees are outside this protected area.
shortest leaves nearly oval, the longer leaves oblong, IUCN: NT
with more or less abruptly tapering base and apex,
flat and thin, acute or often cuspidate; leaf colour
Uses
lustrous dark green; midrib on adaxial (upper) side
prominently raised from base to apex, 1 mm wide, No economic uses are recorded for this species at
830 conspicuous but more flattened on abaxial side. present. Most trees are relatively small and do not
Stomata small and inconspicuous, in numerous in- yield quantities of timber, even though the wood is
termittent lines on abaxial (under) side of leaves. of good quality, to make exploitation worthwhile. It
Pollen cones axillary, sessile, solitary or in groups was undoubtedly used in the past by settlers for light
of 2–3, narrowly cylindrical, 10–30 mm long, 2–3 construction work, but no records have been found
mm diam. at anthesis; microsporophylls imbricate, documenting this. Outside a few botanic gardens in
broadly triangular, acute, bearing to globose pol- Queensland, this species it not known in cultivation
len sacs. Seed cones axillary, solitary on a stout pe- and is strictly tropical in its horticultural require-
duncle ca. 10 mm long, consisting of an axis with ments.
one fertile scale subtended by two small, deciduous
bracts (‘foliola’) with 1–2 fertilized ovules; recepta-
Podocarpus drouynianus F. Muell., Fragm. 4: 86, t.
cles 15–25 mm long, swelling to become flattened
31. 1863–64. Margbensonia drouyniana (F. Muell.)
pyriform, succulent and red when ripe. Seeds when
A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc.
ripe solitary or in pairs (often with unequal devel-
Isp. Prir., Otd. Biol. 103 (1): 60. 1998. Type: Austral-
opment) at oblique distal end of receptacle, 17–22
ia: Western Australia, Tone River, [?] Maxwell s.n.
mm long, 12–15 mm wide, elliptic to ovoid, with
1867 (holotype MEL[?], isotype K).
smooth green epimatium obliquely curved at base,
olive green ripening to purple. Seeds proper 11–14 Podocarpus brownii C. E. Bertrand, Ann. Sci. Nat.
mm long, obliquely ovoid, with a curved apex, dark Bot., sér. 5, 20: 64. 1874.
brown to nearly black, rugose with one side smooth.
Etymology
Distribution
This species was named after Edouard Drouyn de
Australia: NE Queensland (Atherton Tableland, Bel- Lhuys (1805–1881), minister under Napoleon III and
lenden Ker Range). a member of the Institut de France.
TDWG codes: 50 QLD-QU
Vernacular names
Ecology
Emu berry
Podocarpus dispermus occurs in complex mesophyll
vine forest (rain forest), in which it is a small to me-
Description
dium size understorey tree with large leaves. The
­elevation ranges from 57–971 m (GIS data), it is most Shrubs 1–3 m tall, multi-stemmed and sprouting
common between 500–700 m a.s.l. and the soils (of- from the rootstock, forming dense clumps a few
ten sceletal) are derived from basalt or granite. meters across; stems erect. Bark thin, fibrous, green
turning cinnamon brown. Foliage branches terete,
finely grooved or striate, terminating in a conical,
Conservation
2–4 mm long and 1.5–3 mm wide bud with narrowly
The naturally restricted range (EOO = 2670 km2 and triangular, slightly spreading scales. Adult and juve-
AOO = 1826 km2) of this species with specific habi- nile leaves similar, linear, sessile with a broad, de-
tat requirements would justify the category Near current base, 3–9(–13) cm long, 2–4(–5) mm wide,
acuminate; midrib on adaxial (upper) side obtusely a Mediterranean-type climate, but considering its
raised in proximal part, gradually becoming ob- growth habit it could become weedy and invasive.
scure towards apex, prominently raised along the
full length of abaxial side; leaf colour green above,
glaucous beneath. Stomata in intermittent lines ar- Podocarpus ekmanii Urb., Fedde’s Repert. Sp. Nov.
ranged in two conspicious bands on either side of Regni Veg. 18 (1–3): 17. 1922. Type: Cuba: Holguín
abaxial midrib. Pollen cones axillary, solitary or in Prov., Sierra del Cristal, near the Rio Lebisa,
groups of 2–6, with 10–25 mm long peduncles bear- E. L. Ekman 6790 (holotype S).
ing small scale leaves, cylindrical, at anthesis (5–)
10–20(–22) mm long without peduncle, 2–4.5 mm
Etymology
wide; microsporophylls imbricate, small, apiculate, 831
bearing two globose pollen sacs. Seed cones axillary, The species epithet commemmorates E. L. Ekman, a
solitary, on a 10–20 mm long, slender and bare pe- Swedish botanist, who collected plants in the West
duncle, with 1 fertile scale and 1–2 ovules of which Indies in the period 1915–1930.
usually only one develops; receptacles subtended by
2 deciduous, 2–10 mm long bracts (‘foliola’), grow-
Vernacular names
ing to 20–25 mm long and 10–13 mm wide, fleshy
and swollen at maturity, becoming purple-pruinose No common names have been recorded for this spe-
when ripe. Mature seeds ovoid, 12–20 mm long, cies.
8–12 mm diam., covered in a green or purplish green
epimatium, distal part rounded without a crest; seed
Description
proper ovoid, 7–12 mm long.
Shrubs or small trees to 8 m tall; d.b.h. to 30 cm.
Bark thin, exfoliating in small flakes, light brown
Distribution
weathering dark grey. Branches numerous, spread-
SW Western Australia (from Bunbury to Mt. Bark- ing or ascending. Foliage branchlets slender, terete,
er; Darling Ranges). finely grooved between the densely set leaf bases,
TDWG codes: 50 WAU-WA terminating in small, ovoid-globose buds with im-
bricate, broadly ovate, carinate scales, the outer
scales terminating in an acuminate apex. Leaves
Ecology
densely set towards distal end of branchlets, spread-
Podocarpus drouynianus occurs in the understorey ing at wide angles from shoot, oblanceolate to line-
of jarrah and karri (Eucalyptus spp.) forests on sandy ar-oblanceolate, 1.5–2.7 cm long, straight or slightly
soils, where it forms large or small patches by root curved downward, rarely slightly falcale, 3–5 mm
suckering. It is a lowland species and does not occur wide, gradually narrowing to a short petiolate base,
in the mountains of the Stirling Range N of Albany. more abruptly to an acuminate apex, coriaceous,
It regenerates after fire but is probably dependent stiff, lustrous green or greyish green above, whitish
on the microclimate of the forests for its long-time green below. Midrib 0.5 mm wide, slightly raised
survival. However, it can re-establish in disturbed from leaf base on adaxial (upper) side, often fading
forests and persists in managed forests where log- towards apex, more prominent and continuous on
ging creates temporary openings. It is one of the few abaxial side. Stomata very small but conspicuous in
species in the genus with this habit. numeous white lines on either side of abaxial mid-
rib. Pollen cones axillary, not observed. Seed cones
axillary, solitary on short peduncles 1–2 mm long;
Conservation
receptacles a short axis with two fused bracts, swell-
IUCN: LC ing to 10 × 6 mm, becoming succulent and bright
red when ripe, still showing free acute bract apices.
Seeds solitary, including the epimatium 6–7 mm
Uses
long, 3.5–4 mm wide, narrowly ovoid to ovoid with
No economic uses are known of this species. It may a distal ridge culminating in a 1 mm high crest. Seed
have some potential for horticulture in regions with proper not observed.
 de Moa population(s) is situated in a national park
Taxonomic notes
and here collections were made recently. The trees
This species was treated as a synonym of P. angusti­ were regenerating well in that location. At present
folius Griseb. in my World Checklist and Biblio­ no information is available about the status of the
graphy of Conifers (Farjon, 1998, [2001]). Its most two western populations.
obvious distinctive character is its shorter leaves IUCN: NE
(1.5–2.7 cm long in P. ekmanii versus 2.5–5 cm long in
P. angustifolius), but most other characters appear to
Uses
be similar. Podocarpus angustifolius (here including
P. aristulatus), P. buchii, and P. ekmanii are closely No uses have been recorded of this species.
832 similar and presumably related taxa. While P. buchii
is only known from Hispaniola, P. angustifolius and
P. ekmanii occur partly in the same general area of Podocarpus elatus R. Br. ex Endl., Syn. Conif.: 213.
E Cuba and occupy similar forest formations and 1847. Margbensonia elata (R. Br. ex Endl.) A. V. Bo-
habitats. brov & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
Otd. Biol. 103 (1): 60. 1998. Type: Australia: New
South Wales, Paterson River, R. Brown [3117] (lecto-
Distribution
type BM, here designated). Pl. 36, Fig. 263, 264
Caribbean: Cuba (Holguín Province, Sierra del
Cristal, Sierra de Moa; Santiago de Cuba Province,
Etymology
Loma Azul).
TDWG codes: 81 CUB The species epithet means ‘tall’ and refers to the size
of the trees.
Ecology
Vernacular names
Podocarpus ekmanii occurs in the mountains of E
Cuba in sclerophyllous rain forest on serpentine, Brown pine, Plum pine, Yellow pine
at altitudes between 450 m and 1000 m a.s.l. It is a
shrub or at most a small, stunted tree typical of ‘car-
Description
rascales’ which in Iberian Spanish refers to woods
with Holm oak (Quercus ilex), but here merely to Trees to 30 m tall, with erect trunk to 80 cm d.b.h.
‘stony ground’ occupied by sclerophyllous shrubs Bark thin, becoming longitudinally fissured and
and small trees. It has also been found along streams scaly on large trees, slightly fibrous, brown. Branch-
in these mountains, indicating that it is not as es spreading to form a broad, dense crown. Foliage
drought tolerant as some of the other species in this branchlets terete, smooth, finely grooved, terminat-
formation. The habitat conditions in the disjunct ing in small, conical buds 1.5–2.5 mm long and ca.
populations in the Sierra del Cristal and the Loma 1 mm wide at base, with imbricate, mostly appressed,
Azul, where it occurs between 650 m and 1000 m narrowly triangular and acute scales with free or re-
a.s.l. and 500 m and 700 m resp., are similar to those curved apices. Leaves on seedlings 2.5–6 cm long,
in the larger area further east. 6–8 mm wide, on juvenile and mature plants much
larger but variable, on some trees much smaller
than on others, (3–)5–10(–15) cm long, (5–)7–14
Conservation
(–17) mm wide, linear-lanceolate, tapering gradually
This species is rare and has been collected in three or in wider leaves more abrubtly to a short petiolate
disjunct areas; one is in the Sierra del Cristal, anoth- base; apex acute or sometimes obtuse, often cuspi-
er on the Loma Azul and the third, with possibly 2–3 date; midrib prominent but obtuse on adaxial (up-
subpopulations, is located to the east of these in the per) side, varying in width with leaf but up to 1 mm
Sierra de Moa. Both the Sierra del Cristal and Loma wide, more flattened on abaxial side; leaf colour lus-
Azul (“Sierra Azul”) populations are only known trous dark green above, pale green below. Stomata
from Ekman’s herbarium specimens collected in numerous, small, in two bands of intermittent lines
the early part of the 20th century. Part of the Sierra on abaxial side. Pollen cones clustered, axillary with
 2

833

5
4 3

Pl ate 36 . Podocarpus elatus. 1. Habit of tree. 2. Branchlet with leaves and pollen cones. 3. Pollen cones.
4. Branchlet with leaves and seed cones. 5. Seed cone with two seeds.
 2–4 together, sessile, cylindrical, 10–20 mm long, usually deep, alluvial sands and silts and this species
2–3 mm wide; microsporophylls very small, imbri- is often found on river banks. Associated species
cate, broadly triangular, with two basal globose pol- are e.g. Castanospermum australe, Grevillea robusta,
len sacs. Seed cones axillary, solitary on stout, 3–10 and nearer the coast in a type called mixed noto-
mm long peduncles; receptacles subtended by two phyll vine forest Alphitonia excelsa, Cryptocarya
very small bracts (‘foliola’), growing and swelling to triplinervis, Cupaniopsis anacardioides, Diploglottis
15–25(–30) mm long and 15–20 mm thick at the dis- cunninghamii, and the palm Livistona australis, the
tal end, ripening from red to pruinose dark purple latter often along streams.
or blue-black, very succulent. Seeds 1(–2) on distal
part of receptacle, obliquely attached, ovoid-globose
Conservation
834 with a short proximal beak and an obscure distal
crest, 15–20 × 12–15 mm when mature; epimatium IUCN: LC
ripening from dark glaucous green to purplish black
with a whitish bloom. Seed proper not observed.
Uses
The wood of this species is resistant to termites and
Taxonomic notes
marine borers like barnacles, it is highly durable and
Ken Hill in Flora of Australia 48: 556 (1998) indi- strong. It is therefore valued and used for construc-
cated a collection made by Robert Brown with No. tion like jettys and boat pilings in salt water, for boat
3117, with duplicates at BM and K, as syntype. End- construction, packing cases, carpentry, joinery, and
licher (1847) merely referred to Brown as the author wood turning, and also for furniture, especially out-
of the species and “Habitat in Nova Hollandia ori- door furniture. It takes a fine polish and is therefore
entali” but not to his collections of specimens. We used in high quality products that last long. This
do not know what Endlicher, who was based in Vi- species is occasionally planted as a park or street
enna, has seen, as the conifer specimens at W were ornamental tree; in deforested areas turned over to
all destroyed in World War II. Assuming Brown 3117 agriculture mature individuals may have been re-
is a collection of which Endlicher saw a duplicate tained as amenity trees. It is obviously only suitable
at W and that he may have seen other material col- for these purposes in warm temperate to subtropical
lected by Brown along the Hunter River as well, the climates.
BM duplicate is here designated as the lectotype of
Podo­carpus elatus R. Br. ex Endl., with an isolecto-
type at K. Podocarpus elongatus (Aiton) L’Herit. ex Pers., Syn.
Pl. 2 (2): 580. 1807. Taxus elongata Aiton, Hort. Kew.
3: 415. 1789. Type: South Africa: Cape Province,
Distribution
[“Caput Bona Spei”], F. Masson s.n. (lectotype BM).
Australia: New South Wales, Northern Territory,
Queensland.
Etymology
TDWG codes: 50 NSW-NS NTA QLD-QU
The species epithet means ‘elongated’ and refers to
the leaf shape.
Ecology
Podocarpus elatus occurs naturally in coastal and
Vernacular names
subcoastal rainforest in the region of the eastern
Australian coast which receives ample rain and has Breede River yellowwood; Breërivier geelhout (Af-
a warm climate; it becomes rare in drier forest in the rikaans)
north of Queensland. These forests are regionally
known as complex mesophyll vine forests, charac-
Description
terized by trees with medium-size, entire margined
leaves, an abundance of climbing plants (lianas), and Shrubs or small trees usually 3–6 m tall, but on
locally palms. The altitudinal range of this species is occasion attaining 20 m, d.b.h. to 50 cm; bark on
from near sea level to ca. 1000 m a.s.l. The soils are trees thin, exfoliating in narrow strips, light brown
weathering grey. Shrubs spreading wide to 10–12 leaves observed did not have any stomata on the
m, multi-stemmed, regenerating from fires. Foli- adaxial surface of the leaves.
age branchlets assurgent, slender, terete, with fine
grooves, terminating in buds 2–3 mm diam. with
Distribution
oblong, narrowly triangular, 4–6 mm long outer
bud scales with spreading tips. Leaves crowded in South Africa: Eastern and Western Cape Provinces;
upper parts of branchlets, spreading to nearly erect, Malawi; Zambia; Zimbabwe.
(1.5–)2.5–5(–7) cm long, (3–)4–5(–7) mm wide (to 12 TDWG codes: 26 MLW ZAM ZIM 27 CPP-EC CPP-
cm long and 10 mm wide on juvenile plants or vig- WC
orous epicormic shoots), narrowly oblong-elliptic to
linear, gradually tapering to a petiolate base, more 835
Ecology
abruptly to an acute or sometimes obtuse apex; mar-
gins flat to slightly revolute; leaf colour glaucous to Podocarpus elongatus is an uncommon species
greyish green above, dull whitish green below; mid- growing in woodlands in moist sites, usually along
rib on adaxial (upper) side thin, raised in lower half (intermittent) streams and in ravines, or on rocky
and fading towards apex, on abaxial side continu- sites with sparse vegetation. Woodland species in
ous and distinctly raised. Stomata in two bands of these ravines are Cunonia capensis and Olea capen­
many white lines separated by midrib on abaxial sis, among other trees, and sclerophyllous shrubs on
side, only occasionally a few short lines of stomata the drier edges. In the Western Cape its habitat con-
on adaxial side. Pollen cones axillary, solitary or tacts with fire-prone vegetation types such as fynbos
sometimes in clusters of 2–5, sessile or sometimes and as a consequence individual trees are frequently
pedunculate, cylindrical, 15–25 mm long, elongating burnt. Resprouting from the base, they then de-
to 30–40 mm at anthesis, 3–5 mm diam., subtended velop into broad spreading shrubs or bushes, while
by ovate or broadly rounded, scarious, acuminate only individuals that are protected from fire, e.g. by
bud scales; microsporophylls spirally insertend, im- growing in a deep ravine, can develop into monopo-
bricate, outer part triangular, minutely denticulate dial trees of some size. In the NE part of its scattered
or lacerate, each bearing two oblong basal pollen range it is usually a component of moist evergreen
sacs. Seed cones axillary, solitary, on 4–12 mm long, forest and grows more often into a tree.
stout (1 mm diam.) peduncles; receptacles an axis
with two fused bracts, green but swelling to a scar-
Conservation
let, 9–15 mm long and 10–15 mm wide (at distal end)
succulent body. Seeds including the epimatium 7–12 IUCN: LC
mm long, ellipsoid to ovoid with a slightly narrowed
distal end, glaucous green. Seed proper ovoid with
Uses
purple-mottled seed coat.
Due to its small size and usually bushy habit, this
species is not of economic importance as a timber
Taxonomic notes
tree. It is rare in cultivation, but a few cultivars have
The presence of stomata on the adaxial (upper) side been recorded.
of the leaves in “1 to several short rows … in shal-
low longitudinal grooves” was reported by Leistner
in Flora of Southern Africa 1: 40 (1966) and repeated Podocarpus fasciculus de Laub., Blumea 30 (2): 277.
in several compilations on southern African trees. 1985. Type: Taiwan: Taichung Co., Tashue Shan,
My observations of herbarium specimens at K with [“Tai-shu Shan”], D. J. de Laubenfels P 675 (holotype
a powerful Leica MZ6 binocular microscope at 30– L).
40 × magnification did reveal a few short lines of
stomata on that side on only some specimens (e.g. Podocarpus macrophyllus (Thunb.) Sweet var. liu­
Pearson 5328 from near Oliphant’s River). Nearly all kiuensis Warb., Monsunia 1: 192. 1900; Podo­carpus
stomata are confined to the abaxial side and most macrophyllus (Thunb.) Sweet f. grandifolius Pilg., in
Engler, Pflanzenr. IV.5 [18]: 80. 1903.
 Taiwan and apparently P. fasciculus occurs on sev-
Etymology
eral other islands in the Ryukyus, up to the main
The species epithet (Latin fasciculus = cluster or fas- islands of Japan.
cicle) refers to the clustered pollen cones.
Distribution
Vernacular names
Japan: Pacific side of Honshu, Shikoku, Kyushu,
Given in Flora of Taiwan, ed. 2, 1: 561 (1994) in Chi- Nansei-Shoto (Ryukyu Islands); Taiwan.
nese characters. TDWG codes: 38 JAP-HN JAP-KY JAP-SH NNS TAI

836
Description Ecology
Trees to 20 m tall, with erect, terete trunk. Bark not Podocarpus fasciculus is a small to medium size tree
described. Foliage branches terete, grooved, termi- in evergreen, lower montane to montane forests. It is
nating in buds 4–5(–9?) × 2 mm, with lanceolate, often found to grow as an understorey tree, but may
keeled and spreading scales. Leaves linear-lanceo- reach the canopy in lower forest types on the ocean-
late, short petiolate, on juvenile plants up to 16 cm ic islands of the Ryukyus. Its ecology remains poorly
long and 14 mm wide; on adult plants 5–10(–12) cm described in the available literature. In Taiwan, its
long, (4–)6–10 mm wide, thin and easily bending, altitudinal range is given as between 1500 and 2500
straight or more or less falcate, gradually tapering m a.s.l. by S. Y. Lu in Vol. 1 of Rare and Endangered
at both ends, acute or acuminate; midrib prominent Plants in Taiwan (1996). This six volume work in
on both sides, acutely raised adaxially, wider and Chinese gives localities (with Pinyin transcription),
not acute abaxially; leaf colour bright green above a distribution map with altitudinal range, the IUCN
(adaxially), pale green below. Stomata on abaxial rating of conservation status with criteria, selected
(under) side, very small, in numerous irregular lines herbarium specimens, and a set of colour photo-
on either side of midrib. Pollen cones axillary, ses- graphs for each species. It does not describe habitats
sile, in clusters of (1–)2–5, subtended by small (ca. in any detail but these can be estimated from some
2 × 1 mm) scarious, keeled scales, cylindrical, elon- of the photographs (not for P. fasciculus).
gating to 15–25(–30) mm × 2–2.5 mm; microsporo-
phylls spirally arranged, with a triangular, spreading,
Conservation
0.3–0.5 mm long apex and with two conspicuously
protruding, globular pollen sacs. Seed cones solitary This species is treated as endemic to Taiwan in Flora
and axillary on 6–12 mm long, slender peduncles; of Taiwan (ed. 2) 1: 561, pl. 219 (1994), following De
receptacles 9–16 mm long, with two 2–3 mm long, Laubenfels’s original description, but occurs also
obtuse basal bracts (‘foliola’), swelling when ripe to and more widely in southern Japan, including the
a succulent, red ‘fruit’ 10–12 mm thick. Seeds sin- Ryukyu Islands. The Japanese populations were con-
gle at truncate distal end of receptacle and partly sidered a variety of P. macrophyllus (var. liukiuensis
enclosed by it, ovoid-globose, ca. 9 × 7 mm includ- Warb. or forma grandifolius Pilg.) in the past, but are
ing the green epimatium, ripening to purple-green. here included in P. fasciculus. In Taiwan, it was as-
Seed proper ca. 7 × 5 mm, slightly flattened, brown. sessed as Endangered (EN) in Rare and Endangered
Plants in Taiwan 1 (Lu, 1996), with only five localities
known and populations declining under threat from
Taxonomic notes
selective felling for its timber.
De Laubenfels (op. cit.) described P. fasciculus as IUCN: NT
a new species based on his own collections from
Taiwan, with one of these designated as the type.
Uses
Similar plants were described from southern Japan
(Kyushu: Nagasaki; Ryukyu Islands: Iriomote Is- In Taiwan this species has been exploited for its
land) as P. macrophyllus var. liukiuensis Warb. from timber, but as most trees do not attain large size, its
Kyushu, and as P. macrophyllus f. grandifolius Pilg. commercial importance is limited. The wood is used
from the Ryukyu Islands. Iriomote Island is close to for light construction, carpentry, and sometimes
f­ urniture making. It may be planted in city parks foliage branchlets, axillary on short peduncles; re-
as an ornamental, masquerading under the general ceptacles subtended by a 3–4 mm long bract (‘folio­
name P. macrophyllus, to which it is superficially lum’), ca. 6 mm long when still immature, bilobed.
similar. Seeds in mature state not observed.

Distribution
Podocarpus gibbsiae N. E. Gray, J. Arnold Arbor.
39: 429. 1958 [“gibbsii”]. Type: Malaysia: Sabah, Malaysia: Sabah (Mt. Kinabalu).
Ranau District, Mt. Kinabalu N. P., Marai Parai, TDWG codes: 42 BOR-SB
near camp, J. Clemens & M. S. Clemens 32021
(holotype A). 837
Ecology
Podocarpus gibbsiae occurs on mountain ridges at
Etymology
altitudes between 1200 m and 2400 m a.s.l. in mossy
The species epithet commemorates Lilian Suzette forest within the cloud belt on Mt. Kinabalu. It ap-
Gibbs (1870–1925), who collected and studied the pears to be confined to ultramafic soil derived from
flora of Mt. Kinabalu. serpentine and similar rocks. This forest type has an
open canopy up to 20–25 m tall and consists of a
mixture of angiosperms and gymnosperms (mostly
Vernacular names
conifers); common conifers are Phyllocladus hypo­
No common names are recorded for this species. phyllus and Dacrydium gibbsiae. Epiphytes, from
lichens, mosses, and ferns to orchids are numerous.
Description
Conservation
Small to medium sized trees 7–20 m tall. Bark be-
coming scaly on largest stems, brown weathering The endemic population known on Mt. Kinabalu of
grey. Crown in more or less sheltered trees conical, this species occupies an area (AOO) of less than 100
becoming rounded with age, otherwise usually ir- km2 and the trees are restricted to a specific soil type,
regular. Foliage branches spreading to assurgent, which is discontinuous within that area. Most trees
densely leaved distally, forming tufts of foliage; are now within Mt. Kinabalu National Park. Al-
shaded branches with more widely spaced leaves. though occurring within the cloud belt, exceptional
Terminal buds on leading shoots 4–9 mm long, spells of drought and the greatly increased tourism
2–3 mm wide, narrowly conical, with free spread- on the mountain pose a potential fire hazard, to
ing, acuminate outer scales. Leaves on young which the species is not adapted.
plants 3–6(–9) cm long, 4–7(–9) mm wide, linear-­ IUCN: VU (D2)
lanceolate, straight or slightly curved, acute. Leaves
on adult plants (much) smaller 1–3.5(–4.5) cm long,
Uses
3–6(–7) mm wide, elliptical or obovate-oblong to
linear-lanceolate (longer leaves in shaded foliage?), No uses have been recorded for this tree.
coriaceous, spreading to erect and densely crowded,
gradually tapering to a petiolate base; apex obtuse to
rounded, or slightly apiculate from extended mid- Podocarpus glaucus Foxw., Philipp. J. Sci. 2: 258.
rib, rarely acute; margins slightly revolute; midrib 1907. Type: Philippines: Mindoro, Parabatugan,
acutely raised but fading distally on adaxial side, ob- Mt. Halcon, E. D. Merrill 5672 [holotype PNH
tusely raised from base to apex on abaxial (lower) (destroyed?), isotypes K, NY].
side; new leaves flushing pink. Stomata very small
and inconspicuous on abaxial side. Pollen cones
Etymology
axillary, solitary or rarely in pairs, sessile, elongat-
ing at anthesis to ca. 15 mm, 3–4 mm wide; micro­ The species epithet refers to the bluish (glaucous)
sporophylls with acute apex ca. 1 mm long, bearing leaves of the type specimen; not all leaves of all
two large pollen sacs. Seed cones towards ends of plants of this species are so coloured.
 Vernacular names on mountain ridges. Sometimes it descends from
these down into taller forest, where it can become a
nipa (New Guinea)
subcanopy tree. Its altitudinal range is from 1000 to
2800 m a.s.l. Its substrates are acidic to neutral, e.g.
Description
sandstone or karst limestone; near the Danau Paniai
Decumbent or erect shrubs or small to medium size [Wissel] Lakes in New Guinea it occurs on peaty soil
trees to 15 m tall; trunk of trees monopodial, erect, on the slopes and on rocky terrain at the summit of
to 60 cm d.b.h. Bark of trees fibrous, flaky, brown a mountain. It is associated with numerous shrubs,
weathering grey; inner bark pink. Branches spread- among which are species of Gymnostoma (Casuari­
ing, in trees forming a spreading crown. Foliage naceae) and Rhododendron (Ericaceae).
838 branchlets terete, slender, terminating in small buds
with erect or slightly spreading, triangular to lan-
Conservation
ceolate and narrowly acute scales up to 2 mm long.
Leaves on saplings and young trees usually larger IUCN: LC
than on shrubs and sun-exposed branches of mature
trees, to 3.5 cm long and 7 mm wide, elliptic. Leaves
Uses
on shrubs and mature trees often densely crowded,
when sun-exposed 0.8–1.8 cm long, 3–6 mm wide, No uses have been recorded of this species. It is un-
elliptic to obovate, gradually or more abruptly taper- likely to be of much use for its timber and it is not
ing to a short petiolate, slightly twisted base; apex known in cultivation.
often obtuse to abruptly rounded, sometimes api-
culate; margins slightly revolute. Midrib on adaxial
(upper) side 0.2 mm wide, acutely raised, on abaxial Podocarpus globulus de Laub., Blumea 30 (2): 269.
side wider, to 0.5 mm, obtuse. Leaf colour variable; 1985. Type: Malaysia: Sabah, Tawau District, Gu-
upperside dark green or glaucous; underside light nung Silam, D. J. de Laubenfels P 688 (holotype L).
green or grey-green; new leaves flushing red. Sto-
mata very small, numerous on abaxial face. Pollen
Etymology
cones axillary, solitary, cylindrical, 15–20 mm long,
2–2.5 mm wide, with a few rounded scales (from the The species epithet means ‘a little ball’ and refers to
buds) on a 2 mm long peduncle; microsporophylls the foliage buds.
rounded at apex, with two globose pollen sacs at
base. Seed cones axillary, solitary; receptacles with
Vernacular names
two 1.5 mm long basal bracts (‘foliola’), 6–8 mm long
and swollen when ripe, becoming purple. Seeds in- No vernacular names have been recorded for this
cluding the epimatium ovoid-globose, 6–7 × 5 mm, tree.
with a crest at maturity, glaucous green to purplish
green; seed proper not observed.
Description
Trees to 27 m tall, commonly to 15 m, d.b.h. to 45
Distribution
cm; trunk erect, terete. Bark smooth, eventually
Malesia: Admiralty Islands (Manus), Maluku with small, powdery and flaky scales, brown; inner
[Moluccas] (Seram), Philippines (Mindoro), Su- bark soft and fibrous, red-brown. Branches spread-
lawesi; Papuasia: Bismarck Archipelago, New Guin- ing, forming a rounded crown. Foliage branchlets
ea, Solomon Islands. terete, slightly grooved or ridged, terminating in
TDWG codes: 42 MOL PHI SUL 43 BIS NWG-IJ globose compact buds 2–3 mm diam. with imbri-
NWG-PN SOL-SO cate, rounded or apiculate scales with entire mar-
gins. Leaves on juvenile plants (broadly) lanceolate,
to 15 cm long and 25 mm wide, straight, coriaceous,
Ecology
abruptly tapering to an acuminate apex. Leaves on
Podocarpus glaucus is most commonly a decumbent mature plants linear-lanceolate, (2.5–)4–8(–9) cm
or erect shrub occurring in stunted mossy forest long, (7–)9–15 mm wide, straight or slightle curved,
coriaceous, gradually tapering to a petiolate base; carp’ timber; the modest size of most trees would
apex acute-acuminate, obtuse or rounded; midrib indicate that it is not. Where forest is cleared, the
on adaxial side narrow, to 0.8 mm, prominently more valuable trees will be ‘salvaged’ and put to uses
raised from base to apex, on abaxial side wider, to commonly associated with this timber.
1.5 mm, nearly flat. Stomata very small, in numerous
irregular lines on abaxial (under) side. Pollen cones
axillary, solitary or in clusters of 2–3, sessile or sub- Podocarpus glomeratus D. Don, in Lambert,
sessile, subtended by rounded bracts (bud scales), Descr. Pinus 2: [21]. 1824. Type: Peru: [“Chili.
long cylindrical, 25–45 mm long, 3–4 mm wide; Herb: Pavon.”], R. Ruiz & J. A. Pavón y Jiménez s.n.
microsporophylls ca. 1 mm long, with triangular (holotype BM).
apex and two basal pollen sacs. Seed cones axillary, 839
solitary on a 3–5 mm long, stiff peduncle; recepta- Podocarpus cardenasii J. T. Buchholz & N. E. Gray,
cles subtended by 2 minute, 1–2 mm long, early de- J. Arnold Arbor. 29: 142. 1948.
ciduous bracts, slender, ca. 10 mm long, consisting
of two lobes (axes?) each with a distal bract, one of
Etymology
these being fertile, the lobes swelling mostly distally,
ripening to red. Seeds including the epimatium 7–9 The species epithet refers to the clustered pol-
× 5–6 mm, ovoid-globose with a small crest, green len cones, “collected closely together into a head”
turning dark brown. Seed proper not observed. (Stearn, 1983).

Distribution Vernacular names

Malaysia: Sabah (and on the border with Sarawak). No common names have been recorded for this spe-
TDWG codes: 42 BOR-SB BOR-SR cies.

Ecology Description
This species occurs in lower to middle montane Shrubs or stunted trees up to 12 m tall; trunk of trees
rainforest on ridges and summits where the forest to 35 cm d.b.h. Bark smooth, on larger trunks exfo-
is not dominated by dipterocarps, in some locali- liating in small flakes, dark brown weathering black-
ties on ultrabasic soil; altitudinal range 310–1530 m ish grey. Branches spreading or ascending, forming
a.s.l. based on information on herbarium specimen a bushy crown. Foliage branches stiff, ascending or
labels. erect, terete, with fine grooves running down from
slightly raised leaf bases, terminating in subglobose
buds 3–5 mm wide at base, the scales triangular,
Conservation
carinate, with free, acuminate apices, the outer scale
Only known from four disjunct localities, none of sometimes elongated to 7–10 mm. Leaves on juve-
these is in a protected area. The area of occupancy nile plants similar to those on adult plants, spreading
(AOO) is calculated to be less than 500 km2 based to erect, rigidly coriaceous, (1.5–)2–3.5(–5) cm long,
on these four localities and the fact that it is known 2–4(–5) mm wide, linear-lanceolate or sometimes
from only a few herbarium collections, indicating slightly falcate, gradually narrowing to a short peti-
that it is rare. Its altitudinal range places it in lower olate base; margins revolute; apex acuminate and
to middle montane rainforest, which is known to be strongly spinescent; leaf colour greyish green above,
in decline. None of the presently known collections glaucous below. Midrib a continuous groove on
originates from protected areas. adaxial (upper) side, inconspicuous and only raised
IUCN: EN [B2, a, b (iii)] near leaf base on abaxial side. Stomata very small, in
numerous intermittent lines on abaxial side. Pollen
cones axillary, clustered with 4–6 on a stalk to 10 mm
Uses
long, sessile on this stalk, subtended by acuminate
It is not known whether this relatively rare (and bracts of variable length up to 5 mm and with trian-
unknown) species is specifically exploited as ‘podo- gular, carinate bud scales at their base, 4–6 mm long,
 1–1.5 mm wide; microsporophylls with scarious den-
Vernacular names
ticulate margins and obtuse apex, bearing two small,
globose pollen sacs. Seed cones axillary, solitary, on No common names have been recorded for this spe-
short or longer peduncles to 10 mm; receptacles 5–6 cies.
mm long, becoming succulent and red. Seeds in-
cluding the epimatium globose, ca. 5 mm long, with
Description
or without a minute crest. Seed proper not observed.
Spreading or decumbent shrubs to 2 m tall, branches
assurgent to erect. Bark thin, fibrous, light brown to
Distribution
reddish brown. Foliage branchlets terete, rough with
840 Bolivia, Ecuador, Peru (Apurimac). persistent leaf bases, terminating in small, globose
TDWG codes: 83 BOL ECU PER buds with at base a few free but otherwise imbri-
cate, triangular, 1–2 mm long scales. Leaves densely
crowded, remaining on branchlets for several years,
Ecology
oblanceolate (widest above the middle) to linear,
Podocarpus glomeratus is a species of the high An- 0.8–2.2 cm long (longest on juvenile plants), 1.7–2.3
des occurring in high montane to subalpine for- mm wide, curved downward, gradually tapering to a
ests and woodland or scrub, at altitudes from 1800 short, petiolate and decurrent base; margins strong-
m a.s.l. (but usually not below 2500 m) to 3600 m ly revolute; apex obtuse or rounded. Midrib forming
a.s.l. or perhaps higher. Above 3000 m this species is a groove on adaxial (upper) side, prominently el-
dwarfed and shrubby; below this it is a constituent evated but obtuse on abaxial side. Stomata small, in
of cloud forest rich in epiphytes, especially mosses two narrow bands on either side of abaxial midrib.
and lichens. Pollen cones axillary, solitary on a short peduncle
lacking bud scales at base, subtended by 3–5 keeled
Conservation bracts, cylindrical, 8–14 mm long, 2–2.5 mm wide at
anthesis; microsporophylls triangular, with two sub-
IUCN: LC
basal, semi-globose pollen sacs. Seed cones axillary,
solitary on a short peduncle; receptacles formed of a
Uses
short axis with two basally fused but distally spread-
The wood of this species is exploited for the mak- ing bracts of unequal to nearly equal size, swelling
ing of furniture and cabinets, the smaller and more to become one or two distinct, succulent, bright red
crooked trunks and branches for firewood. It is not bodies, each ca. 7 × 4 mm (if one then slightly larg-
known to be in cultivation. er) in case of two only one of which bears the single
seed. Seeds including the epimatium 6–7 mm long,
obliquely ovoid, 3.5–4 mm wide, the green surface
Podocarpus gnidioides Carrière, Traité Gén. Conif., becoming grooved or rugose, terminating in a crest.
ed. 2, 2: 656. 1867. Podocarpus alpinus R. Br. ex Seed proper not observed.
Hook. f. var. arborescens Brongn. & Gris, Bull. Soc.
Bot. France 13: 425. 1866. Type: New Caledonia:
Taxonomic notes
[locality not stated], F. von Mueller 70 (lectotype P).
This species is similar to Podocarpus nivalis from
Podocarpus alpinus R. Br. ex Hook. f. var. caespitosus New Zealand and P. lawrencei from SE Australia
Pancher ex Brongn. & Gris, Bull. Soc. Bot. France 13: including Tasmania and is perhaps most closely re-
425. 1866; Podocarpus gnidioides Carrière var. caes­ lated to the latter. The original description by Car-
pitosus Pancher ex Carrière, Traité Gén. Conif., ed. rière (op. cit.) stated that this species is a tree, with
2, 2: 657. 1867. the shrub as a variety, but this species never grows
into a tree and only shrubs exist. De Laubenfels
(1972) cited “Mueller 70” as the holotype of P. gnidi­
Etymology
oides Carrière and as the type of P. alpinus Hook. f.
The species epithet refers to Gnidia (Thymelaea­ var. arborescens Brongn. & Gris, but these authors
ceae), to which it is supposed to bear a resemblance. merely referred to an unidentified collection by Mu-
eller of “Podocarpus spec.” (Carrière) and “Mueller,
Vernacular names
1862” (referring to the year of collection). More than
one specimen from Mueller in P are thus marked No common names are known for this species.
and none of these authors annotated them with their
taxon names. For these reasons, Mueller 70 at P is to
Description
be considered the lectotype chosen and designated
by De Laubenfels of both names. Trees to 40 m tall, with erect trunk to 1.5 m d.b.h.
Bark thin, becoming scaly on large trees, flak-
ing in narrow strips, slightly fibrous, brown; inner
Distribution
bark pinkish. Branches spreading to form a broad
New Caledonia: Grande Terre, Province Sud. crown. Foliage branchlets terete, smooth, finely 841
TDWG codes: 60 NWC grooved, terminating in large, globose buds 3.5–4.5
mm long and 4–5 mm wide, with imbricate, round-
ed or broadly triangular scales with appressed or
Ecology
sometimes a few free apices. Leaves on seedlings
Podocarpus gnidioides is a shrub of the higher 3–4 cm long, 5 mm wide, on juvenile and mature
mountains (above 600 m a.s.l.) of the southern part plants much larger, (6–)10–25(–30) cm long, 7–18
of New Caledonia (Grande Terre) and has been mm wide, linear, straight or slightly curved, taper-
found close to the summit of Mont Humboldt at ing gradually to a petiolate base; apex acute; midrib
1600 m a.s.l. It forms broad shrubs in rocky terrain slightly raised and obtuse on adaxial (upper) side,
with low vegetation on serpentine rock, where there ca. 1 mm wide, often more prominent on abaxial
is no competion from trees and where it grows with side; leaf colour dark green above, glaucous green
numerous other montane plants, among which are below. Stomata numerous, small, in two bands of
a number of ferns, that are adapted to nutrient poor intermittent lines on abaxial side. Pollen cones axil-
and ultramafic soils. lary, solitary or with 2–4 together, sessile, subtended
by imbricate, rounded scales, cylindrical, 25–45 mm
long, 4–5 mm wide; microsporophylls very small,
Conservation
imbricate, broadly triangular, with two sublateral
IUCN: NT globose pollen sacs. Seed cones axillary, solitary on
slender, 5–10 mm long peduncles; receptacles sub-
tended by two very small, deciduous bracts (‘folio-
Uses
la’), growing and swelling to 9–14 mm long and 7–9
Although this species is very similar in habit and mm thick at distal end, ripening to orange-yellow
foliage to the Australian and New Zealand shrubby then red. Seeds solitary on distal part of recepta-
podocarps, it has not been taken into cultivation cle, obliquely attached, ovoid-globose with a short
beyond a few botanic gardens. It is obviously less proximal beak and an obscure distal crest, including
hardy than the other two, but cultivation of plants is the epimatium 12–15 × 9–12 mm when mature; epi­
not necessarily restricted to countries with frost and matium ripening from glaucous green to dark red.
snow in winter and it is therefore to be recommend- Seed proper not observed.
ed that it be tried for rock gardens in a mild climate.
Distribution
Podocarpus grayae de Laub., Blumea 30 (2): 275.
Australia: N Queensland, Northern Territory (Arn-
1985 [“grayii”]. Type: Australia: Queensland, Cape
hem Land).
York Peninsula, Parrot Creek (Annan River drain-
TDWG codes: 50 QLD-QU NTA
age), L. J. Brass 20203 (holotype L). Fig. 265, 266, 267

Etymology Ecology
This species was named after Netta Elizabeth Gray Podocarpus grayae is widespread in the subcoast-
(1913–1970), who (in part with J. T. Buchholz) pub- al warm subtropical to tropical rainforests of N
lished a serial monograph of the genus. Queensland, where it occurs from near sea level to
 ca. 750 m a.s.l. It can occur on coastal flats directly ing a broad or sometimes pyramidal crown. Foli-
behind the mangroves, along streams, and on low age branchlets terete, finely grooved from decurrent
mountain ridges. In Arnhem Land, Northern Terri- leaf bases. Terminal vegetative buds semi-globose or
tory, a few small, relict populations occur in patches ovoid, ca. 4 mm wide, with imbricate, broadly tri-
of rainforest along streams. angular, keeled, acute, sometimes apiculate scales
2–3 mm long; axillary buds small, with ovate scales.
Leaves on young plants larger than on mature
Conservation
plants, to 16 cm long and 18 mm wide. Leaves on
IUCN: LC mature trees lanceolate to linear-lanceolate, straight
or slightly curved, sometimes falcate, 4–10 cm long,
842 6–14 mm wide (largest on shaded lower branches),
Uses
gradually or more abruptly narrowing to a short
Large trees are a valuable source of timber but data petiolate base; margins parallel or nearly parallel in
on its exploitation are lacking, primarily due to the middle part of leaf, gradually tapering to an acute to
fact that it has been described as a new species rela- slightly acuminate or sometimes obtuse apex. Mid-
tively recently. It has been erroneouly identified as rib continuously raised but narrow, not in a groove,
P. neriifolius in the past and its uses may therefore on adaxial (upper) side, forming a more or less
have been similar to those of that widespread (but prominent and continuous ridge abaxially. Stomata
not Australian) species. It is reported to be in culti- in numerous intermittent lines in broad bands on
vation in some botanic gardens. either side of midrib on abaxial side. Pollen cones
axillary, sessile, solitary, cylindrical, 10–15 mm long;
microsporophylls triangular, acute, with two basal
Podocarpus guatemalensis Standl., Proc. Biol. Soc. pollen sacs. Seed cones axillary, on short peduncles;
Washington 37: 49. 1924. Type: Guatemala: Izabal, receptacles 6–8 mm long, swelling and becoming
Puerto Barrios, P. C. Standley 25090 (holotype US). succulent, reddish turning brown. Seeds includ-
ing the epimatium ellipsoidal, 7–8 mm long, 5 mm
Podocarpus pinetorum Bartlett, Publ. Carnegie Inst. wide, smooth with a small, apical crest, light brown
Washington 461: 21. 1935; Podocarpus guatemalen­ or grey-brown. Seed proper not observed.
sis Standl. var. pinetorum (Bartlett) J. T. Buchholz &
N. E. Gray, J. Arnold Arbor. 29: 137. 1948.
Taxonomic notes
Podocarpus allenii Standl., Ann. Missouri Bot. Gard.
28: 409. 1941; Podocarpus guatemalensis Standl. var. Bartlett (op. cit.) described from Mountain Pine
allenii (Standl.) J. T. Buchholz & N. E. Gray, J. Ar- Ridge, Belize a new species P. pinetorum, compar-
nold Arbor. 29: 137. 1948. ing it with the type of P. guatemalensis, which ap-
parently came from a shrubby young plant with
longer, falcate leaves. He “reluctantly separated [it]
Etymology
from P. guatemalensis” because he realized that
The species epithet indicates Guatemala, the coun- it might just represent the more mature form of
try from which the species was first known. that species. Likewise, Standley (op. cit.) expressed
doubts about the distinction between his new spe-
cies P. ­allenii from Panama and P. guatemalensis on
Vernacular names
similar grounds. Leaves of these large Panamanian
Ocotillo, Ocotillo de llano (Spanish) trees measured only 3.5–4.5 cm long and 7–8 mm
wide, but leaves of seedlings and saplings measured
9–12 cm long and 9–14 mm wide. These two species
Description
were reduced by Gray & Buchholz (1948) to varieties
Trees to 20–35 m tall, erect; trunk up to 1–1.5 m d.b.h. of P. guatemalensis, mainly on the basis of asserted
Bark smooth, reddish brown turning dark grey and differences in leaf anatomy, particularly the presence
fissured on large trees. Branches spreading, form- of sclereids and their position in the ­mesophyll. Orr
(1944) found sclereids only in P. oleifolius among Podocarpus henkelii Stapf ex Dallim. & A. B. Jacks.,
American species of Podocarpus, but he did not Handb. Conif.: 47. 1923 [“henckelii”]. Type: South
examine leaves of these two taxa and only juvenile Africa: Kwazulu Natal, Bulembu Forest, Mt. Ayliff,
leaves of P. guatemalensis. As Gray & Buchholz ad- Fort Donald Forest Station, W. G. Cochrane s.n.
mit, there may be a correlation with leaf form (juve- (lectotype PRE). Fig. 268
nile versus adult) as well as environment (lowland
versus upland), and taxonomic distinction on the Podocarpus ensiculus Melville, Kew Bull. 1954: 566.
basis of these characters in limited samples may not 1955.
be justified.
Etymology
843
Distribution
This species was named after Heinrich Henkel, a
Belize; Colombia; Costa Rica; Ecuador (Cordillera German botanist who died in 1914.
del Condor); Guatemala; Honduras; El Salvador;
Mexico (Chiapas); Panama; Venezuela (Cordillera
Vernacular names
Oriental, Selvas de Guatopo).
TDWG codes: 79 MXT-CI 80 BLZ COS ELS GUA Henkel’s yellowwood, Falcate yellowwood; nanjula
HON PAN 82 VEN 83 CLM ECU (Malawi)

Ecology Description
Podocarpus guatemalensis occurs in mixed coni- Trees to 35 m tall; trunk to 1.8 m d.b.h. Bark thin,
fer-angiosperm forest or pine forest, often along fibrous, smooth but on large trunks fissured, exfolia-
streams. The altitudinal range calculated from data ting in longitudinal strips and small or large flakes,
with herbarium specimens is 1–2150 m a.s.l., but it light brown or tan weathering grey; inner bark pink-
is most common below 1200 m a.s.l. It is therefore a ish red, drying to red-brown. Main branches spread-
species both from lowland and montane forests. In ing or ascending, forming a rounded crown. Foliage
the lowlands it is often present in savanna type veg- branches terete, finely grooved from decurrent leaf
etation with Pinus oocarpa, P. caribaea, and many bases, usually stout, terminating in globose to short
angiosperm shrubs; it there occupies stream sides. conical buds 3–5 mm wide with imbricate, apiculate
In evergreen broad-leaved tropical rainforests it is a inner scales and a few longer outer scales with re-
canopy tree, apparently successfully competing with curved margins and spirally rolled apices. Leaves on
other trees. Little is known of the soil types in which seedlings and saplings to 22 cm long, 7–12 mm wide,
this tree occurs in the latter vegetation type; often linear, straight to falcate. Leaves on mature trees
conifers are restricted to nutrient-poor substrates spreading wide or often drooping to nearly pen-
where they do better with the aid of mycorrhiza. dulous, (5–)8–12(–17) cm long, (5–)7–9(–10) mm
wide, linear, straight or falcate, narrowing to a peti-
olate base and very gradually tapering to an acute
Conservation
or acuminate apex, thin coriaceous; margins slightly
IUCN: LC revolute; midrib indistinct and raised only in proxi-
mal two thirds of leaf on adaxial (upper) side, more
distinct and continuous but narrow on abaxial side;
Uses
leaf colour lustrous dark green above, dull green
No specific data could be found regarding the uses below. Stomata on abaxial side, in numerous inter-
of this species. It will undoubtedly be logged with mittent lines on either side of midrib, small. Pollen
other trees and sold on a limited scale as podocarp cones axillary, solitary or in clusters of 2–5, sessile,
wood, which is used for construction, carpentry, subtended by rounded to ovate, scarious, keeled bud
and, to a lesser extent, depending on size and quality, scales with lacerate margins, subglobose becoming
furniture, veneer, and tool making. It is not known cylindrical, elongating to 4–5 cm, 4–5.5 mm wide;
to be in cultivation. microsporophylls spirally inserted on a stout rachis,
 imbricate, ovate-triangular, each bearing two oblong c­ odominant in Ocotea-Podocarpus forest with a can-
pollen sacs at base. Seed cones axillary, solitary on a opy 30–40 m tall and both Afrocarpus ­usambarensis
5–10 mm long, stout peduncle, initiated as a short and P. milanjianus are often also present as canopy
axis with 2 bracts not or only very weakly enlarging trees.
to a clavate receptacle (hence a succulent, coloured
receptacle is usually absent). Seeds single, enclosed
Conservation
in a thick epimatium, 17–22 × 14–18 mm, broadly el-
lipsoid, green or olive green ripening to yellowish IUCN: LC
or purplish and pruinose, exuding a whitish latex
from distinct vescicles when cut. Seed proper not
Uses
844 observed.
This species is a valuable timber tree but not com-
mon enough to be of major economic importance.
Taxonomic notes
The wood is used for construction, carpentry and
Podocarpus henkelii is unusual in the genus, because joinery, as well as furniture making. In South Africa
its seed cone bracts, one of which is fertile and on and Zimbabwe it is commonly planted as an orna-
which develops the large seed, do not or sometimes mental tree; outside the indigenous region of the
only slightly enlarge and never develop into a large, species it is often planted in southern California. It is
succulent and coloured receptacle. The epimatium also in cultivation in England and Wales and young
is thicker than in other species and colours pur- trees are offered to the trade from British nurseries
plish pruinose (as in a damson or small plum) and (Grimshaw & Bayton, 2009: 641).
presumably takes over the function of attraction
to birds as food. This development and the result-
ing ‘fruit’ resembles that in Afrocarpus, which to- Podocarpus hispaniolensis de Laub., Moscosoa 3:
tally lacks a receptacle. The vegetative characters of 149. 1984. Type: Dominican Republic: Cordillera
P. henkelii are different from Afrocarpus (e.g. hypo­ Central, Peravia, Rancho Arriba, near Finca de
stomatic leaves) and (in toto) identical with other Los Suizos, M. Mejía & J. Pimentel 444 (holotype
species of Podocarpus. In Nageia, commonly lacking JBSD).
a receptacle, one species (N. wallichiana) usually de-
velops a small but succulent and coloured one, but
Etymology
the epimatium of the seed, given its size, colour, and
texture, presumably remains the main food attrac- The species epithet refers to the island of Hispaniola,
tion. Apparently the development of a receptacle in where this species is endemic.
these genera is not a completely fixed character.
Vernacular names
Distribution
No common names have been recorded for this spe-
South Africa: Eastern Cape Province, Kwazulu Na- cies.
tal; Malawi; Tanzania; Zimbabwe.
TDWG codes: 25 TAN 26 ZIM 27 CPP-EC NAT Trees 15–20 m tall; trunk massive, d.b.h. 1–1.5 m in
large specimens. Branches spreading and drooping,
forming a wide, domed crown. Bark not described.
Ecology
Foliage branchlets slender, terete, with fine grooves
Podocarpus henkelii is a tree that occurs in montane between leaf bases, terminating in small, globose
evergreen rainforest, often on steep, rocky slopes, at buds 2–3.5 mm diam. and 2–3 mm long with imbri-
altitudes between 1300 m and 2000 m a.s.l.; it is also cate scales; outer scales keeled, with apiculate, erect
present in coastal forests near sea level in Eastern or incurved apices. Leaves on saplings presumably
Cape Province (where it is rare) and Kwazulu Natal. larger than on mature trees. Leaves on mature trees
Here it may be associated with Afrocarpus falcatus; spreading or somewhat drooping, 3.5–8 cm long,
both are in these forests emergents above a cano- elliptic-linear, 6–11 mm wide, gradually narrow-
py of angiosperm trees. In Tanzania it occurs as a ing to a short, 1–3 mm long petiole and to an acute
or apiculate apex. Midrib on adaxial (upper) side Podocarpus humbertii de Laub., Adansonia 11 (4):
lying in a distinct groove, on abaxial side promi- 714. 1971. Type: Madagascar: Antsiranana Prov.,
nently raised and continuous to apex. Pollen cones Anjanaharibe Massif, Mont Anjanaharibe, slopes
axillary, solitary, sessile, observed only in the im- and north summit, west of Andapa, H. Humbert et
mature state before elongation to full length. Seed al. 24741 (holotype P).
cones axillary, solitary on a 7 mm long peduncle;
receptacles formed of an axis with two fused, fleshy
Etymology
bracts, swelling to 8–10 mm long and 6 mm wide
and becoming succulent and red. Seeds including This species has been named after H. Humbert, who
the epimatium ca. 7 mm long and 4 mm diam. with collected it in 1951.
a distinct distal crest. 845
Vernacular names
Distribution
No common names have been recorded for this spe-
Caribbean: Hispaniola, Dominican Republic (Cor- cies.
dillera Central, Prov. Puerto Plata).
TDWG codes: 81 DOM
Description
Small, shrubby trees 3–15 m tall or perhaps larger.
Ecology
Bark exfoliating in small flakes, brown weathering
This species is found in tropical broad-leaved ever- grey. Foliage branchlets spreading to erect, slen-
green rainforest in the mountains of the Domini- der, terete, with persisting leaf bases, terminating
can Republic at altitudes between 750 m and 1200 in small, subglobose buds with imbricate, triangu-
m a.s.l. It grows with many different trees, one of lar, obtuse scales. Leaves small, crowded towards
which is the palm Prestoea acuminata, which has ends of branches, spreading at 50° or less from
been found with it in several locations. Compared shoot, curved downward towards apex, overlapping
to its moderate height, trees have been found with each other particularly near ends of branchlets and
massive trunks (one was blown by hurricane ‘David’ concealing buds, obovate to oblanceolate (the wid-
in 1984), which may indicate that it is a long-lived est part towards the apex), 7–15 mm long, 2–4 mm
species of which the larger individuals have with- wide, gradually narrowing towards a petiolate base;
stood high winds for a considerable time. margins flat or slightly revolute, abrubtly narrowing
to form an obtuse, rounded or sometimes acumi-
nate apex; texture coriaceous; leaf colour dark green
Conservation
above, dull green below. Midrib narrow, inconspic-
This species is only known from five locations, all uous and discontinuous or in a groove on adaxial
visited for collection of herbarium specimens in (upper) side, continuous and wider but flattened on
the 1980s. Its total area of occupancy (AOO) is esti- abaxial side. Stomata small, in numerous intermit-
mated to be less than 500 km2 and due to deforesta- tent white lines on either side of abaxial midrib. Pol-
tion several populations are fragmented. Logging of len cones axillary, solitary or in pairs on a 3–6 mm
large trees has had an impact on the number of ma- long peduncle, cylindrical, 8–20 mm long, 2.5–3 mm
ture trees, which have become scarce in some areas. wide; microsporophylls with a triangular, ca. 1 mm
IUCN: EN (B1+2e) long apex and two basal, globose pollen sacs. Seed
cones axillary on a short, 2–3 mm long peduncle; re-
ceptacles formed of an axis with two unequal bracts
Uses
and enlarging to only 3–5 × 2.5–4 mm, becom-
This species is being logged together with other ing ­coriaceous. Seeds solitary, including the thick
large trees in the forests where it occurs. The wood epimatium 11–15 mm long, 9–12 mm wide, ovoid-­
is used for general construction and carpentry. It is globose, with a small, inconspicuous crest below the
not known to be in cultivation. apparent apex.
 Distribution Vernacular names
Madagascar: Antsiranana Province. In Papua New Guinea this species is known locally
TDWG codes: 29 MDG as ‘dala’ and ‘tunum’ (Milne Bay); in the Solomon
Islands ‘dengali’ or ‘dingale’ and ‘mou’ were recorded
by collectors.
Ecology
Podocarpus humbertii is reported from sub-humid
Description
forest, dry lowland deciduous forest, and ericoid
thickets or wooded heath on mountain summits of Trees to 25(–35) m tall, to 60 cm d.b.h., bole straight,
846 gneiss and granite. The elevation ranges from 1600 to 20 m. Bark smooth, thin, on large boles with nar-
m to 2410 m a.s.l. according to data on herbarium row, longitudinal flakes or strips, light brown weath-
specimen labels. In the original description of the ering grey; inner bark pinkish or reddish brown,
species it was reported to occur up to 2800 m a.s.l. fibrous. Branches spreading, forming a rounded or
domed crown. Foliage branchlets terete, more or less
grooved, glabrous, terminating in 2–3 mm long buds
Conservation
with spreading, triangular to lanceolate outer scales.
This species is know from five to seven subpopula- Leaves on juvenile plants short petiolate, linear-
tions (locations) and most of the herbarium collec- lanceolate, 10–15 cm long, (7–)12–18 mm wide, of-
tions were made between 1950–1960 with only one ten curved and with undulating margins, gradually
recent collection in 2001. Based on the mapped her- narrowed at base, long acuminate or acute. Leaves
barium collections, the extent of occurrence (EOO) of mature trees shorter or of similar length, shade
is calculated as 2103 km2, which falls within the leaves 8–15 cm long, (7–)12–18 mm wide, acuminate;
threshold of Endangered (EN). Its habitat is (in part) leaves more exposed to light mostly linear-lanceolate,
dry lowland deciduous forest, which is under pres- 5.5–10(–13) cm long, 8–15(–18) mm wide, straight or
sure from grazing. A continuing decline is inferred slightly curved, petiolate at a more abruptly narrow-
from the collection dates and the degradation of ing base, tapering to an acute or obtuse apex. Midrib
parts of its habitat. This species has been collected in acutely raised adaxially, flat or obtusely raised abaxi-
the following protected areas: Tsaratanana Reserve, ally, usually narrower on adaxial side; leaf colour dark
Marojejy National Park, and Anjanaharibe-Sud Spe- green above, pale green below, new leaves flushing
cial Reserve. Only one collection, from Mont Tsara- pale whitish green. Stomata very small, in numerous
tanana, was made recently, in 2001. irregular lines on abaxial (under) side on either side
IUCN: EN [B1 a-b (iii, iv)] of midrib. Pollen cones axillary, solitary or in clus-
ters of 2–3, sessile or short pedunculate, cylindrical,
elongating to 25–40 mm, 2–3 mm wide; microsporo-
Uses
phylls spirally arranged, triangular, spreading, with
No economic uses have been recorded of this spe- two globose pollen sacs. Seed cones axillary, solitary
cies. It is probably used for firewood locally. on a 5–10 mm long peduncle; receptacles a short axis
to 10 mm long with two fused bracts, swelling to be-
come red and succulent when ripe. Seeds solitary
Podocarpus insularis de Laub., Blumea 30 (2): at truncate end of receptacle, enclosed in a smooth
266. 1985. Type: Papua New Guinea: Louisiades epimatium, green turning purplish when ripe, ovoid
Archipelago, Tagula Island [Sudest Island], to subglobose, 10–15 mm long, without a crest. Seed
L. J. Brass 27987 (holotype L). proper not observed.

Etymology Taxonomic notes

The species epithet refers to its insular (on islands) This species appears to be more variable in its leaf
distribution. shapes and size than described by De Laubenfels
(1985, 1988), at least in material seen at the Kew
Herbarium (K) that seems to be best assigned to it.
Unfortunately, most specimens are sterile, and both
Vernacular names
pollen cones and seed cones have been described
here more or less provisionally. No common names have been recorded for this
species other than the generic ‘pinheirinho’ which
means little more than ‘conifer’.
Distribution
Papuasia: New Britain, New Guinea (including Lou-
Description
isiades Archipelago), Solomon Islands; Southwest
Pacific: Vanuatu (Erromango, Tanna). Small trees to 10–12 m tall, d.b.h. to 30 cm. Branches
TDWG codes: 43 BIS NWG-IJ NWG-PN SOL-SO 60 spreading, forming a rounded or irregular crown.
VAN Foliage branchlets slender, terete, longitudinally 847
grooved between leaf bases, terminating in small,
subglobose buds with imbricate, broadly triangu-
Ecology
lar, carinate, short apiculate scales. Leaves on sap-
Podocarpus insularis occurs as a canopy tree in sub- lings similar but slightly larger than on mature trees.
montane to montane rainforests mostly on islands Leaves on mature trees crowded, spreading at wide
in the Bismarck, Solomon, and Vanuatu Archipela- angles or more often at narrow angles to shoot and
gos. It is usually scattered, but locally common and directed forward, (1–)2–4(–5) cm long, (1.5–)2.5–4
attains considerably size in sheltered forests, but re- mm wide, lanceolate-linear to linear, straight or
mains a low, stunted tree on exposed mountain ridg- sometimes slightly falcate, or curved downward,
es. It has been collected from near sea level to 1680 m gradually narrowing to a sessile or short petiolate
a.s.l. It occurs mostly in angiosperm forest (e.g. with base, gradually tapering to an acute or pungent apex;
Nothofagus at higher altitudes in Papua New Guin- margins (nearly) parallel for most of the leaf length,
ea), but can be associated with a few other species of slightly revolute; texture coriaceous or thin coria-
Podocarpus, among which are the similar P. neriifo­ ceous; leaf colour green or grey-green above, whit-
lius and the small-leaved P. glaucus on ridges. ish green below; flushing leaves reddish, subtended
by enlarged perular scales. Midrib ca. 0.4 mm wide,
continuous or sometimes fading towards apex,
Conservation
mostly forming a groove on adaxial (upper) side,
IUCN: LC more or less raised from a concave leaf surface and
continuous to apex on abaxial side. Stomata small, in
numerous intermittent lines forming two bands on
Uses
abaxial side. Pollen cones axillary, on short shoots
Larger specimen trees yield valuable timber, used or slender peduncles up to 15 mm long in 1-several
for light construction, carpentry, boat building clusters of 2–8 or more, cylindrical, 5–12 mm long,
(especially oars), and flooring. This species is usu- 1.5–2 mm wide; microsporophylls ovate-triangular,
ally not recognized as distinct from P. neriifolius by with minutely denticulate margins, bearing two
foresters and loggers and simply taken as podocarp globose pollen sacs. Seed cones axillary, solitary on
wood. It is not known to be in cultivation anywhere. slender, 5–10 mm long peduncles; receptacles small,
swelling to 6–7 mm long, 4–5 mm wide, succulent,
reddish purple. Seeds solitary, globose, including
Podocarpus lambertii Klotzsch ex Endl., Syn.
the epimatium 4–5 × 4–5 mm, glossy without a crest.
Conif.: 211. 1847. Type: Brazil: [locality not stated],
Seed proper globose with a smooth surface.
F. Sello s.n. (holotype W, destroyed, isotypes BM,
K). Pl. 37
Distribution
Etymology
NE Argentina: Missiones; S Brazil: Sao Paulo, Rio de
This species was named after Aylmer Bourke Lam- Janeiro, Paraná, Santa Catarina, Rio Grande do Sul.
bert (1761–1842), compiler and publisher of the TDWG codes: 84 BZL-MG BZL-RJ BZL-SP BZS 85
­famous elephant folio ‘A Description of the Genus AGE-MI
Pinus’.
848

1
7

6
4 2
5

Pl ate 37. Podocarpus lambertii. 1. Habit of tree. 2. Branchlet with leaves and pollen cones. 3. Pollen
cones. 4. Branch with leaves and seed cones. 5. Seed cone. 6. Leaf, upperside. 7. Leaf, underside.
 Toulouse T. 2, 1 (2, 21): 66. 1976 (nom. inval., Art.
Ecology
33.2).
The habitat type most commonly cited for Podo­
carpus lambertii is ‘campo rupestre’ (‘rocky grass-
Etymology
land’) which is in actual fact a mozaic of different
habitats, composed of usually stony fields with The species epithet latifolius means ‘with broad
grasses and often also shrubs, wet or peaty areas, leaves’.
small and larger streams bordered with taller woody
vegetation, or even with gallery forest. In the east-
Vernacular names
ern States where this species occurs, it will be mostly
gallery forest, which enters into the Atlantic Forest Broad-leaved Yellowwood, True Yellowwood; 849
belt, especially in Santa Catarina and Rio Grande do Opregte geelhout (Afrikaans); ‘umGeya’ (Xhosa)
Sul. Here P. lambertii approaches the Atlantic coast
and may also occur, with or without P. sellowii, in
Description
the drier types of Atlantic Forest. Podocarpus lam­
bertii is also found in rocky areas where rain runoff Trees to 30 m tall or more in forests, or stunted small
may provide sufficient moisture. Its small, narrow trees 2–3 m tall on exposed mountain slopes, bole
leaves are adapted to periods of drought. Few data of large trees to 1.2 m d.b.h. Bark thin, smooth, light
are available indicating altitude: some collections brown, exfoliating in long strips, weathering grey.
were made from 900 m to 1800 m a.s.l. Crown relatively small in forest trees, low spread-
ing in stunted trees. Foliage branchlets stout, terete
or slightly angular, finely grooved from the decur-
Conservation
rent leaf bases, terminating in obtuse buds 2–3.5
This species has a large range; the extent of occur- mm diam. with narrowly triangular, acuminate,
rence (EOO) calculated from mapped herbarium sometimes recurved outer scales. Leaves on juvenile
specimens = 761,575 km2. The human footprint plants up to 17 cm long and 20 mm wide. Leaves on
factor is fairly high (45), especially near the coast, mature trees shorter but variable and often crowded
where habitat and individual trees are likely to have towards ends of branchlets, (2–)3.5–8(–12) cm long,
disappeared. The IUCN Conifer Specialist Group mostly straight, (5–)6–12(–18) mm wide, elliptic to
feels it is warranted to flag this species as NT. The linear-elliptic or linear, longer leaves with parallel
species is not known to occur in any protected area. sides in middle part, gradually or more abruptly
IUCN: NT narrowing to a petiolate base, abruptly narrowing to
an obtuse or mucronate to apiculate apex; margins
slightly revolute; leaf colour dark green to glaucous
Uses
green above, dull green below; flushing leaves bright
No commercial uses have been recorded for this green or bronze. Midrib thin and inconspicuous
small tree, of which the timber has not much value. on adaxial (upper) side, fading towards upper part,
It may be used locally for fence posts and most likely more prominent and continuous on abaxial side.
for firewood. It is not known to be in cultivation. Stomata in numerous intermittent lines on either
side of abaxial midrib, rarely a few present on upper
leaf surface. Pollen cones axillary, solitary or rarely
Podocarpus latifolius (Thunb.) R. Br. ex Mirb., in pairs, sessile or on very short peduncles, cylindri-
Mém. Mus. Hist. Nat. 13: 75. 1825. Taxus latifolia cal, elongating to 20–30 mm long at anthesis, 3–4
Thunb., Prodr. Pl. Cap.: 117. 1800. Type: South Af- mm wide, subtended by 3–4 mm long, narrowly tri-
rica: Cape Province, [“Houtniquas, Grootvaders- angular, keeled bud scales; microsporophylls trian-
bosch, aliis”], C. P. Thunberg UPS 23780 (holotype gular to broadly ovate, lacerate, bearing two oblong
UPS). Fig. 270 pollen sacs. Seed cones axillary, solitary on slender
5–15 mm long peduncles; receptacles composed of an
Podocarpus latifolius (Thunb.) R. Br. ex Mirb. var. axis with two bracts, one or sometimes both fertile,
latior Pilg., in Engler, Pflanzenr. IV.5 [18]: 90. 1903; swelling to 8–14 mm long and 8–12 mm wide and
Podocarpus latior (Pilg.) Gaussen, Trav. Lab. Forest. becoming succulent, turning from glaucous green to
 pink and sometimes reddish or bluish purple when plantation, which is unfortunate as that role is most-
ripe. Seeds including the epimatium obovoid to sub- ly taken by several species of Eucalyptus, some of
globose, 7–11 mm long, slightly apiculate or crested, which have turned out to be invasive pests.
glaucous green turning purple or violet. Seed proper
smooth, the seed coat containing resin cavities.
Podocarpus laubenfelsii Tiong, Blumea 29 (2): 523.
1984. Type: Malaysia: Sabah, Ranau District, Mt.
Distribution
Kinabalu N. P., Bukit Burong, D. J. de Laubenfels P
South Africa: from the Cape to the Northern Trans- 715 (holotype L). Fig. 269
vaal.
850 TDWG codes: 27 CPP-EC CPP-WC LES NAT OFS
Etymology
SWZ TVL-GA TVL-MP TVL-NP TVL-NW
This species has been named in honour of David J. de
Laubenfels, a long-time student of Podo­carpaceae.
Ecology
Podocarpus latifolius is a canopy forest tree in the
Vernacular names
coastal and midland primary forests where there is
sufficient rainfall and natural protection from fires No common names have been recorded for this spe-
to allow such forest types to develop. In open coastal cies.
bushland and on dry, rocky mountain slopes it only
grows to a stunted tree a few meters tall at most. It
Description
occurs from near sea level to 2000 m a.s.l. In forested
valleys near the coast it can be associated with Afro­ Trees to 40 m tall, bole erect, straight, to 80 cm
carpus falcatus and both are there commonly emer- d.b.h. Bark nearly smooth, soft, exfoliating in thin
gents above a lower canopy of angiosperm trees, strips, light brown. Branches spreading, in mature
among which members of the families Celastra­ceae, trees forming a rounded crown. Foliage branchlets
Araliaceae and Flacourtiaceae, as well as Olea capen­ terete, glabrous, finely grooved or striated, terminat-
sis are often seen. ing an a small, globose bud up to 4 × 4 mm, with
triangular, imbricate scales forming a short conical
apex, the tips of the scales at apex often recurved.
Conservation
Leaves on saplings and young trees usually larger
IUCN: LC than on mature trees; shaded leaves also larger than
sun-exposed leaves on mature trees, to 25 cm long
and 25 mm wide. Leaves on mature trees exposed
Uses
to sun (7–)10–19 cm long, 10–20(–25) mm wide,
Broad-leaved Yellowwood is a valuable timber tree long petiolate, linear-lanceolate to linear, straight
producing even-grained, light weight, pale yellow or slightly curved, more or less abruptly widening
wood suitable for a variety of purposes. It was ex- above the 6–15 mm long petiole, more gradually
tensively used in colonial times for railway sleepers tapering to an acute or more commonly acumi-
and construction, and many houses were built with nate apex. Midrib on adaxial (upper) side obtusely
it, such as the old Cape homesteads of which many raised, ca. 1 mm wide, on abaxial side wider and flat-
still exist. It is still valued for indoor carpentry and ter, fading towards apex. Stomata very small and in-
floors, as in former times, but large trees have be- distinct, in numerous irregular lines on abaxial side.
come much less common and smaller sizes are now Pollen cones axillary, clustered with 3–5 on a short
used for furniture making, especially when well sea- peduncle or sessile, raising from globular buds with
soned and nicely figured with darker streaks. Afri- rounded scales, cylindrical, elongating to 20–40
cans value the wood for coffins. In South Africa this mm or longer, 2,5–3.5 mm wide; microsporophylls
species is commonly planted as an amenity tree in with an elongated apex, bearing two elongated pol-
parks and along streets. Elsewhere it is uncommon len sacs. Seed cones axillary, solitary on slender pe-
and mainly represented by specimens in botanic duncles (3–)10–15 mm long; receptacles with 2 very
gardens. It is too slow growing for profitable ­forestry short (1–2 mm) bracts (‘foliola’) at base, ca. 10 mm
long. Seeds including the epimatium at least 10 mm making, interior trim, household utensils, and wood
long but fully mature phase not observed. carving. It is not known to be in cultivation.

Distribution
Podocarpus lawrencei Hook. f., London J. Bot. 4:
Borneo: Malaysia: Saba (Mt. Kinabalu), Sarawak 151. 1845., [“lawrencii”] Podocarpus alpinus R. Br.
(Lawas); Indonesia: Kalimantan Timur. ex Hook. f. var. lawrencei (Hook. f.) Hook. f., Fl.
TDWG codes: 42 BOR-KA BOR-SB BOR-SR Tasmania 1 (5): 356. 1857. Type: Australia: Tasmania,
R. W. Lawrence 218 (holotype K). Fig. 271
Ecology
Podocarpus alpinus R. Br. ex Hook. f., London J. Bot. 851
Podocarpus laubenfelsii occurs scattered in ‘keranga’ 4: 150. 1845.
forest with Agathis borneensis, Nageia wallichiana,
Sundacarpus amarus, Dacrydium gracile, and Fal­
Etymology
catifolium falciforme, often on nutrient-poor and/or
water-logged, acidic soils. The species is also scat- This species was named after Robert William Law-
tered in primary rainforest and mossy forest; grow- rence (1807–1833), who collected plants in Tasmania
ing as a large emergent tree on rocky ridges; it may around 1826.
be more common in ‘heath forests’ at higher eleva-
tions. Altitude of P. laubenfelsii ranges from 920 m
Vernacular names
to 1650 m a.s.l. This species occurs with scattered
individuals in the forest, but can be dominant in Mountain plum pine, Plum pine
‘heath forests’ at higher altitudes.
Description
Conservation
Procumbent or more or less erect shrubs to 4 m tall,
Most collections of this species are from Mt Kina- usually creeping and prostrate, forming 4–5 m wide
balu National Park. Deforestation is widespread clumps. Branching often profuse, with densely set
outside of protected areas in the region, indicating foliage branchlets and in creeping shrubs with long
a considerable decline in the past that has not abat- leaders (whip shoots), terminating in small, globose
ed. Outside Mt. Kinabalu National Park it is known buds 1–2 mm diam. with broadly triangular, imbri-
from only three locations, one just outside the park, cate scales. Leaves sessile, decurrent, 4–16 mm long,
the other two at considerable distance. Its total range linear-oblong, the shortest leaves nearly oval, 4–5
is probably incompletely known as the species was mm wide, gradually tapering to a slightly twisted,
only described and named in 1984. It is a large forest narrow base; apex obtuse, sometimes minutely api-
tree with considerable timber value. Its only protect- culate; midrib nearly absent on adaxial (upper) side,
ed location so far known is in Mt. Kinabalu National prominent but obtuse on abaxial side; leaf colour
Park. mid-green or dark green above, light green below
IUCN: EN [B2 a-b (ii)+(iii)] with two grey-green stomatal bands. Stomata small,
in intermittent, wavy lines on either side of abax-
ial midrib. Pollen cones axillary and solitary or in
Uses
groups of 2–3(–6) at distal end of foliage branchlets,
This species attains large sizes in primary lower sessile or on short peduncles, cylindrical, 4–7 mm
montane rainforest and is consequently a valuable long, 2–2.5 mm wide; microsporophylls imbricate,
timber tree logged and traded as other ‘podocarp’ very small, broadly triangular, enclosing towards
trees, without distinction to species or even genus. their base two relatively large, globose pollen sacs.
Its excellent wood is used for house construction and Seed cones axillary, solitary, sessile or shortly pe-
carpentry and for making oars, spars and masts of dunculate; receptacles while growing ca. 3 mm long,
sailing vessels. More specialized uses requiring high swelling strongly at maturity to 5–6 mm, becoming
grade timber are veneer, furniture making, cabinet subglobose, succulent and dark red. Seeds solitary at
distal end of receptacle, narrowly ovoid, 4.5–5 mm
 long, 2.5–3 mm wide, with a narrowing distal end; Podocarpus ledermannii Pilg., Bot. Jahrb. Syst. 54:
epimatium olive green. Seed proper not observed. 210. 1916. Types: Papua New Guinea: West Sepik,
Sepik River, [“Lordberg”], C. L. Ledermann 9943,
C. L. Ledermann 10064a (syntypes K, lectotype not
Distribution
designated).
Australia: New South Wales, A. C. T., Victoria, Tas-
mania. Podocarpus idenburgensis N. E. Gray, J. Arnold Ar-
TDWG codes: 50 NSW-NS TAS VIC bor. 39: 447. 1958.

Ecology Etymology
852
Podocarpus lawrencei is a subalpine to alpine shrub This species was named after the German botanist
limited to the highest mountains in the southern Carl Ludwig Ledermann (1875–1958), who first col-
part of the Great Dividing Range and in Tasmania. lected it in the Sepik River delta.
Its altitudinal range is between 1100 m and 2030 m
a.s.l. and it grows mostly in rocky terrain, e.g. scree
Vernacular names
slopes, broken rocky plateaus and ridges formed by
acidic igneous or metamorphic rock types. In the sua (Papua); babako (Papua New Guinea mainland);
Great Dividing Range it may also occur in wet scle- neleel, nelil (New Britain)
rophyll forest with Acacia spp., Eucalyptus spp., and
Telopea sp., where it can reach to 4 m tall; above the
Description
tree line it is found in subalpine/alpine dwarf scrub
mixed with alpine herbaceous grassland, some asso- Trees to 25(–30) m tall; trunk to 60 cm diam., erect.
ciated species there are Prostanthera cuneata, Grevil­ Bark more or less fibrous, dark brown. Branch-
lea australis and Eucalyptus australis. In Tasmania, es spreading, forming a rounded crown. Foliage
three endemic conifers are often associated with Po­ branchlets terete, glabrous, finely grooved or striate,
docarpus lawrencei: Diselma archeri (Cupressaceae), terminating in an elongated bud 6–12 mm long, 3–4
Microcachrys tetragona, and Pherosphaera hookeri­ mm wide at base, with free, erect to slightly spread-
ana (Podocarpaceae). The angiosperm flora is also ing lanceolate scales drawn out in a caudate apex.
distinct, with e.g. Orites revoluta, Richea scoparia, Leaves lanceolate to linear-lanceolate, 8–20(–22)
Epacris serpyllifolia, and Leptospermum ruprestre. cm long, (11–)15–25 mm wide, straight or slightly
curved, distinctly petiolate, more or less abrubtly
widening at base, with parallel sides for most of
Conservation
their length, gradually tapering to an acuminate or
IUCN: LC sometimes acute apex. Midrib distinctly raised and
narrow on adaxial (upper) side, wider and flattened
on abaxial side; leaf colour lustrous green on both
Uses
sides. Stomata very small, in numerous irregular
Mountain plum pine is suitable as a low, hardy shrub lines on abaxial side on either side of midrib. Pol-
in countries with mildly cold winters. It is planted in len cones axillary, on a 3–4 mm long peduncle, in
rockeries or as undergrowth in park-like tree plant- groups of 1–3, subtended by 4–5 mm long acuminate
ings and provides an evergreen, spreading shrub bracts (bud scales), elongating to 3.5–4.5 cm, 4 mm
with attractive, red ‘fruits’ (the ripe receptacles) wide at anthesis; microsporophylls apiculate, bear-
topped by a shiny, green seed. It is uncommon in ing two elongated pollen sacs at base. Seed cones
Europe, North America and Japan probably because axillary, solitary, on a 5–15 mm long peduncle; re-
it is not easy to germinate; in Australia and New ceptacles subtended by 2 recurved, 2 mm long bracts
Zealand gardeners seem to have a preference for (‘foliola’) and with one apical bract visible when still
conifers from the northern hemisphere, too. growing, at maturity 10–16 mm long, ripening to a
swollen, succulent, orange then red imitation fruit.
Seeds including the epimatium ovoid, 10–13 × 8–10
mm, with a distal, barely elevated crest.
 large boles with narrow, longitudinal flakes or strips,
Distribution
light brown weathering grey; inner bark pinkish or
Papuasia: New Britain, New Guinea. reddish brown, fibrous. Branches spreading, form-
TDWG codes: 43 BIS NWG-IJ NWG-PN ing a rounded crown in large trees. Foliage branch-
lets slender, terete, finely grooved between remote
leaf bases. Terminal buds 3–9 mm long, outer scales
Ecology
lanceolate, terminating in an acuminate, slightly
Podocarpus ledermannii occurs as a scattered and lo- spreading apex. Leaves on saplings and young trees
cally common tree in tropical evergreen rainforests, larger than on mature trees, 12–22 cm long, ca. 15
from near sea level to at least 1500 m a.s.l. It has been mm wide, acute, obtuse or truncated at apex. Leaves
found in forests in river deltas (e.g. Purari, Sepik) as on mature trees linear-lanceolate, (5–)6–11(–14) cm 853
well as in upland lower montane forests dominated long, (7–)10–14 mm wide, straight or slightly falcate,
by Castanopsis. It often grows in the understorey, narrowing at base to a 4–9 mm long petiole; margins
but can attain canopy height in medium-high forests often somewhat wavy; apex acute, obtuse or nar-
or in more open (disturbed?) forests. rowly rounded. Midrib on adaxial (upper) side ob-
tusely raised, especially near base, mostly less than
Conservation 0.5 mm wide and often fading towards apex, raised
and continuous on abaxial side, 0.5–1 mm wide. Sto-
IUCN: LC
mata small, in numerous intermittent lines in two
bands on either side of abaxial midrib. Pollen cones
Uses
axillary, solitary or in clusters of 2–3, sessile, with
This species is relatively uncommon and is probably several basal, carinate bud scales, cylindrical, elon-
not distinguished from the more common species gating to 50–60(–80?) mm, 2–3.5 mm wide; micro-
like P. neriifolius. Therefore it can be assumed to be sporophylls triangular, spreading, with two globose
logged when occurring in accessible forest and at- pollen sacs. Seed cones axillary, solitary on a short
taining suitable size. The uses of the wood are similar peduncle; receptacles subtended by two 2–4 mm
to that of P. neriifolius, i.e. primarily light construc- long bracts (‘foliola’), consisting of an axis with 1–2
tion and carpentry, and in the delta areas where it fertile bracts and 1–2 sterile ones, fusing and swell-
occurs boats, oars, masts, and spars. As far as known ing, initially 5–6 mm long, becoming ca. 8 × 6 mm
this species is not in cultivation. and red when ripe. Seeds single, sometimes two on a
receptacle, including the epimatium 8–11(–13?) mm
long, 7–8 mm wide, ovoid-globose with a rounded
Podocarpus levis de Laub., Blumea 24 (2): 496. distal end. Seed proper not observed.
1979. Type: Indonesia: Papua, Yapen Island,
Mariatu, L. J. van Dijk bb 30484 (holotype L).
Taxonomic notes
This species is rather similar to P. neriifolius, from
Etymology
which it is distinct in its often bluntly rounded leaf
The species epithet means ‘with a smooth surface’; it apices. That character is, however, variable even on
refers to the upper surface of the leaves from which the same branch, with truncate and acute leaf apices
the midrib scarcely protrudes. e.g. on C. B. Robinson 309 (K) from Amboina, which
was cited in the protologue of this species, but the
variation was not commented upon. Podocarpus levis
Vernacular names
does not have acuminate leaves, as seen in juvenile
marisa, sanru (Sulawesi); wasiwarare (Yapen Island, plants of P. neriifolius, and the midrib on the adaxial
New Guinea) side is only prominent near the leaf base and soon
flattens or even fades out towards the apex (at least in
sicco). Whether to recognize such leaf differences as
Description
character states of a distinct species is a moot point;
Trees to 25(–35?) tall, usually with a straight bole; we need more and better material to decide, and this
trunk d.b.h. 40 cm or more. Bark smooth, thin, on species is here merely given the benefit of doubt.
 bases; terminating in obtuse-conical buds with sev-
Distribution
eral elongated, spreading to recurved outer scales
Malesia: Borneo (Kalimantan Timur), Sulawesi, 4–10 mm long. Leaves on juvenile plants (includ-
Maluku [Moluccas]; Papuasia: New Guinea (Papua). ing seedlings) similar to leaves on mature plants but
TDWG codes: 42 BOR-KA MOL SUL 43 NWG-IJ sometimes longer, to 14 cm. Leaves on mature trees
linear-lanceolate, 5–10(–13) cm long, 5–10(–12) mm
wide, straight or slightly falcate, gradually widening
Ecology
from a petiolate base with parallel sides for most of
Podocarpus levis is a scattered tree, locally common, their length; apex acute or sometimes obtuse. Mid-
in evergreen primary rainforest. It has been found to rib on adaxial (upper) side conspicuous but obtuse,
854 grow from near sea level to 1650 m a.s.l. It occurs on sometimes obscure towards apex, on abaxial side
various substrates; in Borneo (one locality known) it flattened but well visible. Leaf colour lustrous green
was found on limestone. above, light dull green below. Stomata very small,
forming numerous intermittent lines in two bands
on abaxial side. Pollen cones axillary, nearly sessile
Conservation
or short pedunculate, solitary or with 2–3 together,
IUCN: LC subtended by short, triangular bud scales, cylindri-
cal, elongating unequally to 15–25 mm, 2.5–3.5 mm
wide; microsporophylls triangular, with an obtuse
Uses
apex and bearing two partly hidden, subglobose
This species is a valuable timber tree and logged pollen sacs. Seed cones axillary, solitary on slender
with other species where large trees occur and are peduncles 8–17 mm long; receptacles subtended by
accessible. The wood is used for general construc- two spreading bracts (‘foliola’) 2.5–3.5 mm long, con-
tion, carpentry and furniture making. This species sisting of an axis with 3–4 bracts, 2 often fertile and
is not known to be in cultivation. spreading distally, swelling and amalgamating to an
8 × 4–5 mm large, succulent and red body. Seeds
1–2, rarely 3 on distal part of receptacle, including
Podocarpus longifoliolatus Pilg., in Engler, the covering epimatium 7–9 mm long, obliquely
Pflanzenr. IV.5 [18]: 79. 1903 [“longefoliolatus”]. ovoid, with a small distal crest fading at maturity.
Type: New Caledonia: Grande Terre, Province Sud, Seed proper not observed.
Mt. Mou, J. A. I. Pancher s.n., 1870 (holotype P).
Distribution
Etymology
New Caledonia: Grande Terre (Province Sud, Île des
The species epithet refers to the two (not very long!) Pins).
bracts at the base of the growing receptacle of the TDWG codes: 60 NWC
seed cone.
Ecology
Vernacular names
Podocarpus longifoliolatus has been found on a few
No common names are recorded for this species. summits above 1100 m a.s.l. in the southern part of
Grande Terre, the main island of New Caledonia.
It occurs on serpentine rock in montane rainforest
Description
with both angiosperms and conifers, the latter dom-
Trees to 20 m tall with a monopodial trunk to 60 inated by Araucaria spp. Rainfall is high and cloud
cm d.b.h. Bark longitudinally striated or fissured, cover or mist frequent throughout the year. The rar-
fibrous, exfoliating in strips, brown or reddish ity of this species, although real, is in part artificial
brown, weathering grey. Branches spreading, form- as it is difficult to find or even recognize. More re-
ing a rounded or irregular crown. Foliage branch- cently (1982) it was found on the Île des Pins at ca.
lets terete, more or less ridged from decurrent leaf 180 m a.s.l. in a moist ravine on Mt. N’ga.
 or purple. Seeds including the epimatium ovoid-
Conservation
globose, 7 × 4–5 mm, with a distinct crest at matu-
This species appears to occur in montane forests at rity, glaucous green to purplish green; seed proper
or near the summits of mountains; some are in pro- not observed.
tected reserves. Due to deforestation its few popula-
tions are severely fragmented.
Distribution
IUCN: EN [B1ab(iii)+2ab(iii)]
Philippines: Luzon (Mt. Tapulao), Mindoro (Mt.
Halcon).
Uses
TDWG codes: 42 PHI
No uses have been recorded of this species. 855
Ecology
Podocarpus lophatus de Laub., Kalikasan 7 (2): Podocarpus lophatus is a species of mountain sum-
137. 1978. Type: Philippines: Luzon, Zambales mits and upper slopes, where it occurs in ‘heath’,
Mountains, Mt. Tapulao, H. M. Curran & scrubland and upper montane, low canopy mossy
M. L. Merritt PFB 9511 (holotype US). forest. Its altitudinal range is from 1800 m to 2585 m
(the summit of Mt. Halcon) a.s.l. On Mt. Halcon it is
very common; its abundance in the type locality at
Etymology
Mt. Tapulao is not mentioned.
The species epithet is derived from Greek (prefix in
compound words: lopho-) and means crested; it re-
Conservation
fers to the seed.
Although no immediate threats are putting this spe-
cies in danger of extinction, the fact that it is still
Vernacular names
only known from two separate mountains in the
No common names are recorded for this species. Philippines, at one of which at least it was said to
be abundant in 1995, puts it under the D criterion as
within a threatened category. This species remains
Description
poorly known and may well occur on other moun-
Shrubs or small trees to 6 m tall. Bark not described tains; it may not have been recognized as distinct
or observed. Branches ascending or spreading, from e.g. P. pilgeri, which has similar leaves and can
forming a dense crown. Foliage branchlets as- be a shrub as well as a large tree.
cending, terete, slender, terminating in small buds IUCN: VU (D2)
3 mm long and 2 mm wide, usually hidden among
crowded leaves. Leaves on shrubs and mature trees
Uses
densely crowded, (1.2–)1.5–3(–3.5) cm long, 4–5(–6)
mm wide, elliptic to oblong, greatest width usually No uses have been recorded of this species. It may
at the middle, gradually tapering to a short petiolate, have some merit for horticulture, but is not known
slightly twisted base; apex acute or nearly obtuse; to be in cultivation.
margins slightly revolute. Midrib on adaxial (upper)
side 0.2–0.3 mm wide, acutely raised, on abaxial side
wider, to 1 mm, obtuse or nearly flat. Leaf colour on Podocarpus lucienii de Laub., Brittonia 12 (1):
upperside dark glossy green, underside light green 80. 1960. Type: New Caledonia: Grande Terre,
or dull green. Stomata very small, numerous, in two Province Sud, Rivière Bleue, D. J. de Laubenfels
broad bands on abaxial side. Pollen cones axillary, P 137 (holotype GH).
solitary, sessile, becoming cylindrical (details of ma-
ture cones not observed). Seed cones axillary to up-
Etymology
permost leaves, solitary; receptacles with two 1.5–2
mm long basal bracts (‘foliola’), 7–10 mm long and This species was dedicated to René Lucien, an ama-
swollen when ripe, glaucous, becoming bright red teur botanist in New Caledonia.
 Vernacular names Distribution
No common names are known for this species. SW Pacific: New Caledonia.
TDWG codes: 60 NWC
Description
Ecology
Small trees to 15 m tall and 30 cm d.b.h. Bark fibrous,
fissured longitudinally and exfoliating in long strips, Podocarpus lucienii is a scattered small tree in moist
yellowish brown weathering grey; inner bark reddish forest occurring at middle elevations (altitudinal
brown. Branches spreading to form a broad crown, range of specimens seen 250–1350 m a.s.l.) in more
856 spreading out wide. Foliage branchlets terete, finely or less open situations on ridges and rocky slopes. In
grooved or ridged, terminating in small, globose or the south it is restricted to serpentine and its ultra-
ovoid buds with 1–2 mm long, appressed and obtuse mafic derivatives, but two localities in the north, on
scales. Leaves on juvenile plants sometimes larger the eastern slopes of Mont Panié and on the western
than on mature plants, to 18 cm long and 20 mm slope of the Roches d’Ouaième, are on micaschist.
wide. Leaves on mature trees and shrubs variable
in length, (3–)4.5–14 cm long, (9–)11–18 mm wide, Conservation
oval-linear (widest at midpoint) to linear, tapering
IUCN: LC
abruptly down to a more or less petiolate base and
to an obtuse or rounded apex, coriaceous, straight
Uses
or only slightly curved. Midrib obtusely raised or
fading to a groove on adaxial (upper) side, 1–1.4 No uses have been recorded of this species.
mm wide, flat or forming a groove in dried leaves
on abaxial side. Leaf colour lustrous grey-green
above, dull light green below. Stomata very small, Podocarpus macrocarpus de Laub., Kalikasan 7
in two bands forming numerous intermittent lines (2): 140. 1978. Type: Philippines: Luzon, Benguet
on abaxial side. Pollen cones from clustered buds in Province, H. M. Curran PFB 10894 (holotype US).
axils of leaves or below these, sessile, solitary or 2–3
together, subtended by small, rounded, imbricate
Etymology
bud scales, cylindrical, 10–20 mm long, 3 mm wide,
yellow turning brown; microsporophylls imbricate, The species epithet refers to the large seed (Latin
peltate, minutely apiculate, bearing two basal, glo- macro = large; Greek carpos = fruit).
bose pollen sacs. Seed cones axillary and solitary
on slender, 12–20 mm long peduncles; receptacles Vernacular names
subtended by two small bracts (‘foliola’), growing to
Malakawayan (Philippines).
9–11 mm long, swelling to 5–6 mm thick, maturing
to red or red-brown. Seeds including the epimatium
Description
relatively large, 13–15 × 9–10 mm, obliquely ovoid
without a crest, smooth, olive green turning brown. Trees to 20 m tall, d.b.h. to 30 cm. Bark peeling in
Seed proper ovoid, smooth, ca. 10 × 7 mm. fibrous strips, brown weathering grey; inner bark
soft fibrous, pink. Branches spreading, forming an
irregular crown in most trees. Foliage branchlets
Taxonomic notes
terete, slightly grooved or ridged, terminating in 2–4
Specimens at K belonging to this species were origi- mm long and 2–4 mm wide, conical or subglobose
nally named P. sylvestris and the two species are buds with triangular to lanceolate, erect to spread-
indeed similar. The leaves of P. lucienii are wider ing outer scales. Leaves on juvenile plants linear to
and terminate more abrubtly at both ends even in linear-lanceolate, 9–15 cm long, 10–14 mm wide,
leaves on juvenile plants; the seeds are also larger gradually tapering to a petiolate base and with an
than those of P. sylvestris, and the receptacles are on acute apex. Leaves on mature trees linear to linear-
longer peduncles. lanceolate, (4–)6–10 cm long, 8–13 mm wide, short
petiolate, abruptly or more gradually narrowed at
base, tapering to an acute or obtuse apex, straight
Conservation
or slightly curved, coriaceous; margins flat or nearly
so; midrib obtusely raised adaxially (above), only IUCN: LC
slightly raised and wider abaxially (below); leaf col-
our lustrous green above, dull and pale green below.
Uses
Stomata on abaxial side, very small and in numer-
ous irregular lines in two broad bands divided by In the Philippines, the wood of this species is used in
midrib. Pollen cones axillary, clustered with (2–)3(– the construction of small aeroplanes as a substitute
4) together, sessile, with a few small scales at base, for Sitka spruce; in musical instruments for sound-
cylindrical, to 25 mm long and 3 mm wide; micro- ing boards, for tennis rackets, and for pencils. It is
sporophylls imbricate, slightly spreading at anthesis, not known to be in cultivation. 857
apiculate, with two oblong pollen sacs. Seed cones
axillary, solitary on a 5–10 mm long peduncle; re-
ceptacles with 2 very small basal bracts (‘foliola’) to Podocarpus macrophyllus (Thunb.) Sweet, Hort.
1.5 mm long (deciduous), 10–15 mm long, swelling Sub. Londin.: 211. 1818.
to 10 mm thick, pruinose, becoming orange-red to
red when ripe. Seeds including the epimatium 14–17
Etymology
× 10–13 mm, globose-ovoid, without a crest, glau-
cous green. Seed proper not observed. The species epithet means ‘with large leaves’; under
Taxus as originally placed by Thunberg (see below)
the leaves would indeed be large.
Taxonomic notes
Although De Laubenfels (op. cit.) mentions some
Vernacular names
distinctions between Podocarpus macrocarpus and
P. rumphii, and in Flora Malesiana ser. 1, 10 (3) (1988) Buddhist pine; kusamaki (Japanese); luo han song
even places these two species in different sections, (Chinese)
they appear to be rather similar. The bud scales of
his own specimens of P. macrocarpus (e.g. P 721) are
Description
not spreading and some seeds in P. rumphii are of
similar size to those of P. macrocarpus. The leaves Trees (or shrubs) to 20 m tall; trunk to 60 cm d.b.h.
of mature trees of P. rumphii are longer than those Bark smooth, peeling off in thin flakes, grey-brown.
of P. macrocarpus. In the key to the species given by Branches spreading or ascending, foliage branches
De Laubenfels (op. cit.) pollen cones were said to be in mature plants dense, forming a spreading crown;
in groups of three, yet they were “in clusters of up to ultimate branchlets angular, glabrous or rarely pu-
at least four” in Flora Malesiana, which is similar to bescent, terminating in a bud with elongated, cau-
the grouping in P. rumphii. Both species are known date and recurved scales. Leaves spirally arranged,
from Luzon, albeit cited from different localities, sessile to short petiolate, leaf blade linear-lanceolate,
with P. macrocarpus more to the north. The latter slightly falcate-lanceolate or (oblong-)oblanceolate,
should perhaps be considered a variety or subspe- (2.5–)4–10(–14) cm long, (3–)4–10(–13) mm wide,
cies of P. rumphii. gradually tapering to base, in juvenile leaves with an
acute apex, in adult leaves with an acute or acumi-
nate apex or more abruptly converging to an obtuse
Distribution
or apiculate apex; midvein acutely raised adaxially,
Philippines: N Luzon. obtusely raised abaxially; leaf colour lustrous dark
TDWG codes: 42 PHI green above (adaxially), pale yellowish or whitish
green below. Stomata very small, in numerous irreg-
ular lines on abaxial side on either side of midvein.
Ecology
Pollen cones axillary, sessile, in clusters of (2–)3–5
Podocarpus macrocarpus occurs as a scattered tree per axil, subtended by several scarious, more or
in tropical montane, evergreen cloud forest (mossy less triangular bracts, long cylindrical, 20–40(–50)
forest) at 2000–2100 m a.s.l. mm long, 2.5–3 mm wide; microsporophylls spirally
 a­ rranged, imbricate, spreading at anthesis, the lami- a low shrub. Many forests at lower elevations have
nar part cordate, with two basal pollen sacs. Seed been cut over and have a secondary vegetation of
cones axillary, solitary, on slender, 5–15 mm long pe- remnant forest trees and pioneer species. In this
duncles; receptacles subtended by 2 recurved bracts vegetation and in remnants of undisturbed forest
(‘foliola’), columnar and glaucous when unripe, 10– Podocarpus macrophyllus occurs with Cunning­
15 mm long, 2–3 mm thick, swelling to 10–12 mm hamia lanceolata, Pinus spp., Podocarpus neriifolius,
thick and becoming yellow, then red or purplish red Cephalotaxus fortunei, Fokienia hodginsii, Taxus
to purple and succulent when ripe. Seeds enclosed chinensis, and where not logged, Pseudotsuga sinen­
in an epimatium, subglobose, 10–12 × 8–9 mm, sis and Tsuga chinensis, as well as numerous angio­
from green ripening to purplish black, sometimes sperms, among which genera in the Fagaceae like
858 pruinose. Seed proper ovoid, ca. 10 × 8 mm, with a Castanopsis and Quercus are conspicuous. Espe-
smooth surface, light brown. cially Podocarpus macrophyllus var. maki has been
widely planted in China and Japan and has probably
been spread by birds into vegetation in which it may
Taxonomic notes
not have occurred previously.
A number of varieties have been described and
were recognized in Flora of China 4: 83–84 (1999),
Uses
but of these only var. maki has here been retained,
while var. chingii is recognized as a distinct spe- Podocarpus macrophyllus (especially var. maki) is
cies P. chingianus. The only difference which seems widely planted in China and Japan as well as in parts
more or less consistent between P. macrophyllus var. of SE Asia as an amenity tree. In Japan, this species
macro­phyllus and var. maki is the leaf width; other is planted for hedges; var. maki is a popular pot plant
measurements, while statistically perhaps signifi- and is occasionally seen as bonsai. A few cultivars,
cant (not tested), show much overlapping. The re- one a compact shrub, others also shrubs but with
ceptacles of living P. macrophyllus var. maki seen in narrow or variegated leaves, are known. The spe-
Taiwan were ripening to red, not purple; those of cies and its var. maki are also commonly planted
var. macrophyllus may turn to purple. in other E Asian countries and find use as an orna-
mental as far away as Puerto Rico. The species was
introduced from China in England in 1804; from
Distribution
Japan (var. maki) in the Netherlands in 1830. Appar-
China: Anhui, Chongqing, Fujian, Guangxi, Gui­ ent shrub forms which occur at higher altitude in
zhou, Hong-Kong, Hubei, Hunan, Jiangsu, Jiangxi, Yunnan would be suitable as ornamentals in large
Sichuan, Yunnan, Zhejiang; Japan; Myanmar [Bur- rock gardens; George Forrest collected such a plant
ma]; Taiwan. from the ranges near Dali in 1906 (Forrest 4665), but
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- this provenance appears not to be in cultivation in
SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS- ­Europe at the present time.
HK CHS-HN CHS-JX CHS-ZJ 38 JAP-HN JAP-KY
JAP-SH TAI 41 MYA 2 varieties are recognized:

Podocarpus macrophyllus (Thunb.) Sweet var.

Ecology
macrophyllus. Taxus macrophylla Thunb., in
Podocarpus macrophyllus is a forest species but also Murray, Linn. Syst. Veg., ed. 14: 895. Mai-Jun 1784;
occurs in secondary vegetation (thickets and scrub Margbensonia macrophylla (Thunb.) A. V. Bobrov
on mountain slopes and hill sides), often along & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
streams. It is one of numerous species in a mixed Biol. 103 (1): 60. 1998. Type not designated. Fig. 272,
mesophytic forest formation, largely constituted of 273
deciduous angiosperm trees and shrubs. Podocarpus
macrophyllus occurs from near sea level to around Podocarpus macrophyllus (Thunb.) Sweet var. pilira­
1000 m a.s.l.; in Yunnan it was recorded (G. Forrest mulus Z. X. Chen & Z. Q. Li, Bull. Bot. Res. North-
4665) growing at 2400 m and higher (–3000 m?) as East. Forest. Inst. 9 (3): 69. 1989.
 HN CHS-JX CHS-ZJ 38 JAP-HN JAP-KY JAP-SH TAI
Description
41 MYA
Adult leaves (3–)5–10(–11) cm long, (5–)7–10(–13)
mm wide, abruptly converging to an obtuse or api- Conservation
culate apex, sometimes subacute.
IUCN: LC

Distribution
Podocarpus madagascariensis Baker, J. Linn. Soc.,
China: Anhui, Chongqing, Fujian, Guangxi, Bot. 21: 447. 1885.
Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan,
Yunnan, Zhejiang; Japan: Honshu, Shikoku; Taiwan 859
Etymology
(extreme S and Lanyu or Orchid Island).
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- The species epithet refers to Madagascar, where this
SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS- species is endemic.
HN CHS-JX CHS-ZJ 38 JAP-HN JAP-SH TAI
Vernacular names
Conservation
Hetatra, Tsindrodravina (Malagasy)
IUCN: LC
Description
Podocarpus macrophyllus (Thunb.) Sweet var. maki
Siebold & Zucc., Abh. Math.-Phys. Cl. Königl. Trees to 25 m tall, but often much smaller and shrub-
Bayer. Akad. Wiss. 4 (3): 232. 1846. Podocarpus by; trunk to 60 cm d.b.h. Bark thin, exfoliating in
macrophyllus (Thunb.) Sweet subsp. maki (Siebold strips on larger trunks, in small flakes on small trees,
& Zucc.) Pilg., in Engler, Pflanzenr. IV.5 [18]: 80. light brown weathering grey. Branches numerous,
1903; Podocarpus maki (Siebold & Zucc.) Gaussen, spreading; foliage branchlets spreading or ascending,
Trav. Lab. Forest. Toulouse T. 2, 1 (2, 21): 214. 1976 stout, terete, finely ridged and grooved, terminating
(nom. inval., Art. 33.2); Margbensonia maki (Siebold in robust, subglobose buds 4–6 mm wide, with im-
& Zucc. ex Endl.) A. V. Bobrov & Melikyan, Bjull. bricate, rounded to ovate scales; lower scales carinate
Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 60. and weakly acuminate, with scarious upper margin.
1998. Type not designated. Leaves highly variable in size, on the type specimen
3–6 cm long, 5–7 mm wide, but ranging from 2–18 cm
Juniperus chinensis Roxb., Fl. Ind. 3: 840. 1832, non long and 3–16 mm wide on other specimens, elliptic-
L. (1767). Podocarpus chinensis Wall. ex J. Forbes, Pi- oblong to linear, mostly thick coriaceous, rigid, but in
netum Woburn.: 212. 1839. one variety more lax and drooping, tapering towards
Podocarpus chinensis Wall. ex J. Forbes var. wardii de a petiolate base and to an acute, acuminate (or long
Laub. & Silba, Phytologia 65: 331. 1988. attenuate) or obtuse apex; margins slightly revolute;
leaf colour lustrous green above, dull green below.
Midrib inconspicuous or obtuse on adaxial (upper)
Description
side, continuous or petering out distally, more distinct
Adult leaves (2.5–)3.5–7(–8) cm long, 4–7(–8) mm and continuous, with abruptly raised edges, on abax-
wide; apex obtuse, acute or acuminate. ial (lower) side. Stomata in numerous intermittent
lines on either side of midrib on abaxial side. Pollen
cones axillary, solitary or with 2–3 together on short
Distribution
peduncles, subtended by imbricate, rounded bud
China: Anhui, Chongqing, Fujian, Guangdong, scales, cylindrical, elongating to 1.5–2.5(–3) cm long,
Guangxi, Guizhou, Hong-Kong, Hunan, Jiangxi, 4–5 mm wide; microsporophylls imbricate, with tri-
Yunnan, Zhejiang; N Myanmar [Burma]; S Japan; angular, minutely denticulate apex, each bearing two
Taiwan. basal, oblong pollen sacs. Seed cones axillary, solitary
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- on 15–23 mm long, slender peduncles, consisting of
YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS-HK CHS- an axis with 2–3 fused bracts which retain free bract
 tips at distal widest end, only swelling slightly to a wood was “extensively used in house building, for
4–5 mm long receptacle (sometimes virtually ab- flooring, etc.” in the latter decades of the 19th century.
sent). Seeds large, ovoid-globose or nearly globose, These uses have diminished with many of the larger
including the epimatium 15–23 mm long , smooth, and more accessible trees now logged. The species
weakly crested, glaucous at first, olive green, ripening and its varieties are not recorded in cultivation.
to brownish black. Seed proper not observed.
3 varieties are recognized:
Taxonomic notes
Podocarpus madagascariensis Baker var. madagas­
Baker (op. cit.) cited three collections as seen by cariensis. Type: Madagascar: [“central Madagas-
860 him in the protologue of this species: R. Baron 2794, car”], G. W. Parker s.n., Aug 1880 (lectotype K, here
R. Baron 3129 and G. W. Parker s.n. All three are rep- designated).
resented by specimens in the Kew Herbarium (K)
and are therefore syntypes. However, whereas the
Description
two collections made by Baron are sterile foliage rep-
resenting young trees (or a single tree), the Parker Leaves coriaceous, more or less rigid and spread-
specimen is an adult foliage branch and has a seed. ing, (2–)3–15(–18) cm long, (3–)5–14(–16) mm wide;
This latter specimen also bears a note from R. Baron apex acute or obtuse. Seeds including the epimatium
(in litt. July 1884) about the uses of the wood and the ovoid-globose.
native name of the tree, which Baker also cites in the
protologue. It is therefore appropriate to designate
Distribution
the specimen at K collected by G. W. Parker as the
lectotype of Podocarpus madagascariensis Baker. Madagascar, along the eastern plateaux and moun-
tains.
TDWG codes: 29 MDG
Distribution
Madagascar, along eastern plateaus and mountains.
Conservation
TDWG codes: 29 MDG
IUCN: LC
Ecology
Podocarpus madagascariensis Baker var. procerus
Podocarpus madagascariensis occurs in moist forests de Laub., Adansonia 11 (4): 715. 1971. Type: Mada-
or forest remnants from level plains near sea level gascar: Toliara Prov., Fort Dauphin, Forêt de Be-
to forested ridges and escarpments at 2000–2400 mangidy, R. Capuron SF 11774 (holotype P).
m a.s.l. In the lowland forest it can attain 20–25 m
and become a canopy tree with mostly evergreen
Description
tropical angiosperms; on exposed rocky ridges it is a
shrub or a stunted tree occurring in sclerophyllous Leaves mostly lax and drooping, linear, 5–13 cm
low forest or scrubland rich in lichens, often on rela- long, 4–6.5 mm wide, thin coriaceous; margins rev-
tively dry sites. The drooping leaved variety procerus olute; apex acute or sometimes obtuse.
occurs from near sea level to about 1000 m a.s.l. on
sand or sandstone; the small leaved var. rotundus oc-
Distribution
curs at altitudes between 1500 m and 1800 m a.s.l. on
igneous rock outcrops. The variety madagascarien­ Madagascar: near Tolanaro [Fort Dauphin] and
sis is much more widespread and occurs on diverse Massif de Bekolosy.
substrates, usually above 800 m a.s.l. TDWG codes: 29 MDG

Uses Conservation
According to one of the collectors of the original This variety was listed as Endangered in a pre-1994
material of this species, the Reverend R. Baron, its IUCN Categories and Criteria listing on the grounds
of its rarity and restricted distribution near Tolanaro
Vernacular names
[Fort Dauphin] in the SE of Madagascar. However, it
is also known from the Massif de Bekolosy in the far ’Cinqui-masé’ (Peru)
north of the island, and could therefore turn up else-
where. Its status as Endangered therefore remains
Description
provisional.
IUCN: EN [E] Trees to 30 m tall; trunk erect, d.b.h. to 60 cm. Bark
of larger trunks becoming fissured, with longitudi-
Podocarpus madagascariensis Baker var. rotundus nally exfoliating strips 3–5 cm wide, dark brown; in-
L. Laurent, Ann. Fac. Sci. Marseille 23: 59. 1915. ner bark pink. Branches spreading to form a broad,
Type: Madagascar: Massif du Manongarivo, 1500– rounded crown in mature trees. Foliage branches 861
1800 m alt., J. M. H. A. Perrier de la Bathie (P, not robust, terete, finely grooved between leaf bases, ter-
designated). minating in subglobose buds 3–5 mm wide at base,
with imbricate, broadly triangular to ovate, carinate
scales with a short acuminate or obtuse apex; axil-
Description
lary buds smaller, globose. Leaves variable in size
Leaves (3–)5–7 cm long, 4–6 mm wide, acuminate especially on different trees, generally largest on sap-
or with a long attenuate apex. Seeds including the lings and young trees, commonly 7–13 cm long and
epimatium nearly globular. 15–20 mm wide, but as small as 3 cm long and 12 mm
wide and as large as 27–29 cm long and 27–30 mm
wide, on saplings acuminate, on mature trees acute,
Taxonomic notes
acuminate or sometimes more or less obtuse in a few
Only known from the Massif de Bekolosy (Bekolosy leaves, lanceolate to nearly linear from a short peti-
River) and Massif du Manongarivo (river) in the olate base, straight or slightly falcate, lustrous green
far NW of Madagascar, but taxonomically perhaps above, dull pale green below. Midrib on adaxial (up-
doubtfully distinct and in need of critical revision. per) side slightly raised, with a groove in the centre,
on abaxial side obtusely raised, to 1 mm wide, con-
tinuous to apex on both sides. Stomata very small, in
Distribution
numerous intermittent lines on either side of abaxial
Madagascar: Massif de Bekolosy, Massif du Manon- midrib. Pollen cones axillary, solitary, sessile, at base
garivo. with spreading perular scales with scarious margins,
TDWG codes: 29 MDG cylindrical, elongating to 30–40 mm, 3–4 mm wide;
microsporophylls triangular, with two relatively
large, oval pollen sacs at base. Seed cones axillary,
Conservation
solitary on 10–20 mm long peduncles; receptacles
A conservation assessment of this variety is diffi- formed of an axis with two unequal bracts, exposing
cult, because very few collections are known in the acute bract tips after swelling, 8–12 mm long, 6–7
herbaria, and its taxonomic distinction is somewhat mm wide at distal end, green ripening to succulent
doubtful. and bright red. Seeds single on the largest bract,
IUCN: DD including the epimatium 9–11 mm long, 5–6 mm
wide, obliquely ovoid, with a crest. Seed proper not
­observed.
Podocarpus magnifolius J. T. Buchholz &
N. E. Gray, J. Arnold Arbor. 29: 133. 1948. Type:
Distribution
Venezuela: Bolivar, La Gran Sabana, Ptari-tepui,
J. A. Steyermark 59989 (holotype F). Central America: Panama; South America: Bolivia
(La Paz), Colombia, Peru (Huánuco, Pasco, San
Martín), Venezuela (Cordillera do Merida).
Etymology
TDWG codes: 80 PAN 82 VEN 83 BOL CLM PER
The species epithet describes the large leaves.
 Podocarpus matudae Lundell, Phytologia 1 (6): 212.
Ecology
1937, [“matudai”]. Type: Mexico: Chiapas, Cerro
Podocarpus magnifolius is a species of tropical mon- Pasitar, E. Matuda 698 (holotype MICH). Fig. 274
tane, evergreen rainforests at altitudes from 850 m
a.s.l. to ca. 1800 m a.s.l. and of cloud forests to an Podocarpus reichei J. T. Buchholz & N. E. Gray,
altitude of 2900 m a.s.l. at least in the Cordillera do J. Arnold Arbor. 29: 131. 1948; Podocarpus matudae
Merida of Venezuela. Not much information has Lundell var. reichei (J. T. Buchholz & N. E. Gray) de
been recorded with herbarium collections, so we Laub. & Silba, Phytologia 68: 69. 1990.
don’t yet know with what other tree species P. mag­ Podocarpus matudae Lundell var. macrocarpus
nifolius is commonly associated. The extraordinary J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 132.
862 large leaves encountered in some trees point to a 1948.
shade tolerant tree that can hold its own with most Podocarpus matudae Lundell var. jaliscanus de Laub.
large-leaved angiosperms and its maximum height & Silba, Phytologia 68: 68. 1990.
would certainly take it into the forest canopy.
Etymology
Conservation
This species was named after its discoverer E. Ma-
Conservation assessment of this (and other) species tuda.
of Podocarpus is hampered by the fact that we do not
know which species are involved in logging opera-
Vernacular names
tions of several that occur within the large range of
P. magnifolius. Based upon that range, and assum- sabino (parts of Mexico).
ing that it is more common than the fairly limited
number of herbarium collections suggest, it might
Description
be considered of Least Concern (LC), since exten-
sive forests still remain in the region. On the other Large trees to 30 m or taller, up to 1.5 m d.b.h.; trunk
hand, the decline may be more serious. The species monopodial, erect. Bark thin, smooth in young trees,
was recognized as distinct in 1948 and for many eventually becoming scaly, light brown weathering
years it remained only known from Bolivia and Ven- grey. Branches spreading or ascending, forming a
ezuela, now we know it from Colombia, Peru and domed crown in large trees. Foliage branchlets slen-
Panama as well. der, straight, terete, with fine grooves from decur-
IUCN: LC rent leaf bases. Terminal vegetative buds 8–12 mm
long, with more or less spreading, long acuminate
scales; axillary buds smaller, 3–4 mm long, with acu-
Uses
minate scales. Leaves of saplings larger than of ma-
This species is widespread but uncommon in the ture trees, to 20 cm long and 17 mm wide; leaves on
forests of Panama and NW South America, where mature trees narrowly lanceolate, straight or slightly
it is undoubtedly logged when encountered for its falcate, flat, 5–14 cm long, 8–13 mm wide (smallest
valuable timber. The wood is used in construction on canopy branches of old, slow growing trees), co-
and carpentry, and perhaps also in furniture mak- riaceous, gradually narrowing to a short petiolate
ing. This species is not known in cultivation, but base, very gradually tapering to an acute or more or
would be of interest in botanic gardens to show to less acuminate apex. Midrib prominent but narrow
the public that conifers can have really large leaves, (less than 1 mm wide) at least in proximal half of
like broad-leaved trees (angiosperms). Its leaf size is adaxial (upper) side, sometimes lying in a shallow
perhaps only surpassed among conifers by the very groove or depression, fading towards apex, thin and
rare species Nageia maxima from Borneo. inconspicuous, sometimes grooved in the middle (in
sicco) on abaxial side. Stomata small and in numer-
ous intermittent lines on abaxial side. Pollen cones
axillary, solitary, sessile, cylindrical, expanding to
30–45 mm long, 4–5 mm wide when full-grown; mi-
crosporophylls with two globose pollen sacs. Seed
cones axillary, solitary on short peduncles; recepta- (800–)1100 m to 2370 m. a.s.l. Most trees in the
cles 6–12 mm long, swelling and becoming succu- broad-leaved forests are oaks (Quercus spp.) but
lent, red turning purplish brown. Seeds including also abundant are other deciduous trees such as
the epimatium 8–12 mm long, broadly ovoid, with Liquidambar, Magnolia, Ostrea, Clethra, and, espe-
an inconspicuous crest, drying dark brown. Seed cially in Chiapas, Puebla and Veracruz, species of
proper ovoid, 6–8 mm long, slightly flattened. northern genera like Fagus, Carpinus and Platanus.
It grows often in ravines near streams. Precipitation
is high, with annual rainfall of 1500–3000 mm and
Taxonomic notes
frequent fog at high altitudes.
Lundell (op. cit.) briefly described P. matudae as a
new species from Chiapas, Mexico, based on E. Ma­ 863
Conservation
tuda 698 with rather short and broad leaves. No
mention was made of the vegetative buds, while ap- The range of this species is extensive, based on the
parently the seed cones were not present in Matuda’s mapping of herbarium collections the extent of oc-
specimens seen by Lundell. Buchholz and Gray (op. currence (EOO) exceeds 600,000 km2, but its dis-
cit.) added a new species from south-central Mexico, tribution is apparently disjunct and scattered. It
P. reichei, but also cited this new species from a col- has been found in a variety of forest types, most of
lection made in Costa Rica. The leaves are similar, if which are under pressure from logging, deforesta-
we ignore their observations about presence or ab- tion, habitat degradation etc., but to what extent this
sence of sclereids in the mesophyll as an informative affects P. matudae is not known. This species occurs
character, but the seed cones (seeds) are larger than in the following protected areas: El Trunfo Reserve,
those in P. matudae. However, Buchholz & Gray also Reserva de la Biósfera El Cielo, Cuenca Hidro­grafica
described, from Chiapas and Guatemala, a new va- del Rio Necaxa and the Pico Pijol Reserves in Mexi-
riety P. matudae var. macrocarpus, with (maximum) co and Honduras.
seed dimensions exceeding those of P. reichei and IUCN: NT
with 4–6 cm long pollen cones. Maturity of these
organs is crucial for proper comparison, as often
Uses
they are not full grown on the specimens examined.
In all this material the buds with their long acumi- The wood of this tree is fine grained, yellow, and
nate scales are most characteristic and I consider it of high quality for building and construction pur-
to belong to a single species occurring from central poses. Podocarpus matudae is slow growing and can
Mexico to Costa Rica. De Laubenfels & Silba (op. only be harvested sustainably at very long rotations,
cit.) described a variety jaliscanus with leaves up to while successful regeneration requires a forest habi-
20 cm long from Jalisco, Mexico, but no information tat with a mixture of other trees (microclimate) as
is given about the range of sizes found in leaves from well as the animal vector for its dispersal. It is there-
this area or whether these long leaves occurred only fore not suitable for plantation forestry. In warm
on vigorous shoots of shaded plants or not. temperate to subtropical countries it would be a
suitable amenity tree for streets and parks. It is occa-
sionally seen in cultivation, mainly restricted to ar-
Distribution
boreta and other living plant collections, and in the
Mexico: Chiapas, Jalisco, Puebla, Tamaulipas, Vera­ western USA usually under its synonym P. reichei.
cruz; El Salvador; Guatemala (Huehuetenango);
Honduras.
Podocarpus micropedunculatus de Laub., Blumea
TDWG codes: 79 MXC-PU MXE-TA MXG-VC MXS-
30 (2): 268. 1985. Type: Malaysia: Sarawak, 4th
JA MXT-CI 80 GUA ELS HON
Division, Marudi F. R., D. J. de Laubenfels P 564
(holotype A).
Ecology
Etymology
Podocarpus matudae is found in mixed pine for-
est, pine-oak forest, montane rain forest, and ever- The species epithet refers to the very short peduncle
green cloud forest, with an altitudinal range from of the seed cone.
 beaches, and in peat-swamp forest, from sea level to
Vernacular names
around 500 m a.s.l. Sometimes this species is asso-
kayu china (Sarawak); kayu tjina (Sabah) ciated with Dryobalanops rappa and Shorea albida
(both Dipterocarpaceae), again as an understorey
tree. This species shares the rare habit of rhizo­
Description
matous propagation with Podocarpus drouynianus
Shrubs or small trees 1–7(–13) m tall, stems 5–20 cm in SW Australia; the latter however never develops
diam., vegetatively reproducing from rhizomes. Fo- into a tree.
liage branchlets terete, glabrous, finely grooved or
striate, terminating in an elongated bud 6–15 mm
Conservation
864 long, 2–3 mm wide at base, with free, erect to slight-
ly spreading linear-lanceolate scales drawn out in a This species is insufficiently known and since its de-
caudate apex. Leaves lanceolate to linear-lanceolate, scription and naming in 1985 few collections have
(6–)8–14(–18) cm long, 10–15(–20) mm wide [some- been made, so that an assessment using herbarium
what larger and wider in juvenile plants than in adult based data was not possible. Its growth habit and oc-
plants, but the distinction is vague due to vegetative currence on the edges of forest clearings seem to in-
reproduction] petiolate, more or less abrubtly wid- dicate that it is probably not in danger of extinction.
ening at base, with parallel sides for most of their IUCN: DD
length, gradually tapering to an acuminate or some-
times acute apex. Midrib acutely raised and nar-
Uses
row on adaxial (upper) side, wider and almost flat
to sunken (in sicco) on abaxial side. Stomata very No uses have been recorded for this shrubby species.
small, in numerous irregular lines on abaxial side on It could perhaps be suitable as a shrub in tropical to
either side of midrib. Pollen cones axillary, sessile or subtropical gardens.
on a very short peduncle, solitary or in groups of
2–3, subtended by 4–5 mm long acuminate bracts
(bud scales), elongating to 35–75 mm, 3–4 mm wide Podocarpus milanjianus Rendle, Trans. Linn. Soc.
at anthesis; microsporophylls apiculate, bearing two London, Bot., ser. 2, 4: 61. 1894. Type: Malawi: Mu-
elongated pollen sacs at base. Seed cones axillary, lanje District, Mulanje Highlands, Mt. Mulanje,
solitary, on a 1 mm long peduncle; receptacles sub- A. Whyte s.n., Oct 1891 (lectotype BM, here desig-
tended by 2 recurved, 3–4 mm long bracts (‘foliola’), nated). Fig. 275
8–10 mm long, ripening to a swollen, succulent, red
‘fruit’. Seeds including the epimatium ovoid, 8–10 ×
Etymology
6–7 mm, with a distal, curved crest.
The species epithet refers to Mt. Mulanje in Malawi,
where Alexander Whyte collected it in 1891.
Distribution
Malaysia: NW Sabah, N Sarawak, Brunei. Vernacular names
TDWG codes: 42 BOR-BR BOR-SB BOR-SR
lusamina (Swahili)

Ecology Description
Podocarpus micropedunculatus is known to occur Trees to 35 m tall; trunk to 1.8(–3.5) m d.b.h., but
as an understorey tree in Agathis forest, where it small and stunted at highest altitudes. Bark thin,
is accompanied by other podocarpaceous conifers smooth or shallowly fissured longitudinally, exfolia-
and by angiosperms adapted to poor sandy soil ting with small, thin flakes or strips, light or dark
(‘kerangas’ forest). It is more abundant and may brown, weathering grey. Branches spreading, form-
form a major element in the vegetation in shrubby ing a broadly domed crown in large trees. Foliage
thickets along the margins of forest clearings, again branches terete, finely grooved or ridged, in slow
mostly on sandy, podzolic soils, or on sandstone. It growing (exposed) crowns with old leaf markings,
also occurs in peaty soils, usually podzolic, of raised terminating in 3–4 mm wide, obtuse buds with a
few spreading and recurving outer scales. Leaves on Rwanda, S Sudan, Tanzania, Uganda, Zaire, Zambia,
seedlings and young trees similar to those on mature Zimbabwe.
trees, but variable in size mostly in relation to expo- TDWG codes: 22 NGA 23 BUR CMN CON RWA ZAI
sure and/or altitude, 1.5–5 cm long on small, stunted 24 SUD 25 KEN TAN UGA 26 ANG MLW MOZ ZAM
trees above 3000 m altitude, 5–17 cm long on trees ZIM
in forests at lower altitudes, 5–16 mm wide, elliptic,
lanceolate-linear to linear, straight or slightly falcate,
Ecology
coriaceous; margins flat or slightly revolute, gradu-
ally tapering down to a petiolate base and to an acute Podocarpus milanjianus is a montane to high mon-
apex. Midrib raised at least from leaf base to beyond tane species occurring in tropical evergreen rainfor-
the middle, situated in a shallow groove, ca. 0.5 mm est, cloud forest, or at its highest limit in dwarf forest 865
wide on the adaxial (upper) side; continuous to apex, dominated by Ericaceae, interfacing with subalpine
wider and flattened on abaxial side. Leaf colour lus- grassland. The altitudinal range is (900–)1300–
trous green above, dull green or sometimes glaucous 3000(–3250) m a.s.l. and about as great in west tropi-
green below. Stomata small, in numerous intermit- cal as in east tropical Africa. It does not form conifer
tent and wavy lines on either side of abaxial mid- forest, although it can occur in forest dominated by
rib. Pollen cones axillary, solitary, sessile or nearly Juniperus procera in e.g. Kenya, and is usually ac-
sessile, subtended by imbricate, broadly rounded, companied by angiosperm trees; however it will of-
carinate bud scales, cylindrical, elongating to 30–40 ten become more abundant on exposed, thin-soiled
mm, 3–3.5 mm wide at anthesis; microsporophylls mountain ridges where the forest becomes low and,
spirally arranged, imbricate, with slightly elongated at high altitudes, even stunted. In tall forest it is
apex and minutely erose-denticulate margin, bear- often restricted to stream sides in deep ravines. In
ing two basal pollen sacs. Seed cones axillary, soli- eastern Africa common associated angiosperm tree
tary on slender ca. 10 mm long peduncles, the two genera in the forests are Albizia, Croton, Macaranga,
bracts (‘foliola’) at base small and early deciduous; Ocotea, Olea, Schefflera, and Syzygium; in Cam-
receptacles consisting of an axis with two bracts fus- eroon Agauria, Albizia, Ilex, Nuxia, Olea, Pyseum,
ing and swelling to a 10–13 × 7–9 mm, succulent red Schefflera, and Syzygium are common with P. milan­
body. Seeds normally solitary at oblique distal end jianus, e.g. on Mt. Oku. ‘Elfin’ forest occurs at the
of receptacle, including the epimatium ca. 8 × 10 highest altitudes above ca. 2800 m and can con-
mm, ovoid-globose without a distal crest, green or sist largely of Erica arborea in eastern Africa, or is
whitish pruinose, turning dark purple. Seed proper mixed, with e.g. Agauria, Cussonia, Myrica, Myrsine,
ca. 6 × 8 mm with a smooth, hard seed coat, brown. Philippia, Syzygium, and Tecomaria. Interfaces with
grassland (Arundinaria alpina) occur here, often
with poor drainage and P. milanjianus concentrated
Taxonomic notes
along rocky streams.
According to the protologue, the type should be
from original material cited as “Habitat in Milan-
Conservation
ji, 6000 ft, Oct. [1891] Nos. 34 & 39” of Alexander
Whyte’s collections, which are kept at BM. In that IUCN: LC
herbarium, the only original material now extant is
a specimen collected by Whyte in October 1891 on
Uses
Mt. Mulanje, but without a number mentioned on
the original sheet fragment; it was separated from This species is an important timber tree in many
another person’s collection and remounted, stating parts of tropical Africa. Its wood is valued for car-
it is “part of Type”. It is here designated as the lecto- pentry and joinery as it is light coloured, even
type of Podocarpus milanjianus. grained, easily worked, and large trees yield good
sizes of sawn timber. More specialized uses requir-
ing high grade timber are veneer, furniture making,
Distribution
cabinet making, interior trim, household utensils,
Tropical Africa: Angola, Burundi, Cameroon, Con- and wood carving. It has been used in afforestation
go Republic, Kenya, Malawi, Mozambique, Nigeria, on a small scale in several African countries, ­within
 and perhaps without its natural range. It is not more or less linear bracts (‘foliola’), 10–14 mm long,
known to be used in horticulture and is probably swelling to 10 mm thick, succulent, changing colour
restricted to a few botanic gardens in Africa and/or from glaucous green to yellow to crimson red when
other tropical countries or in glasshouses in cooler ripe, subtended by two small, leaf-like, deciduous
regions. Provenances from high altitudes may well bracts. Seeds solitary, obliquely ovoid, 8–9 × 6 mm,
be hardy enough to survive in European and other slightly flattened, covered by a pruinose epimatium,
countries with mild winters and abundant rainfall, ripening blackish blue, crested distally. Seed proper
but no one seems to have tried it. ovoid, 7 × 5 mm.

Taxonomic notes
866 Podocarpus nakaii Hayata, Icon. Pl. Formos. 6: 66.
1916. Podocarpus macrophyllus (Thunb.) Sweet var. Podocarpus nakaii is similar to P. macrophyllus and
nakaii (Hayata) H. L. Li & Keng, Taiwania 5: 39. has been treated as a variety of it. The pollen cones
1954. Type: Taiwan: Nantou Co., Holisha, Ta-shu- are solitary or with 2–3 together, not in clusters of
ku, [“Toshoko”], S. Nakai s.n. (holotype TI). Fig. 3–5 as in P. macrophyllus, and the leaves are usu-
276, 277 ally wider, but the ranges of these measurements
overlap. Podocarpus macrophyllus var. macrophyllus
is cultivated in Taiwan, but its var. maki occurs in
Etymology
natural forests at low to middle elevations along the
This species was named after the Japanese plant col- steep eastern coast of the island.
lector S. Nakai.
Distribution
Vernacular names
Taiwan (Chianghua Co.?, Nantou Co., Taichung
A vernacular name has been given in Flora of Tai- Co.).
wan, ed 2, 1: 565 (1994) in Chinese characters. TDWG codes: 38 TAI

Description Ecology
Trees to 20 m tall, with erect, terete trunk. Bark not Podocarpus nakai occurs scattered in broad-leaved
described or observed. Foliage branchlets terete, (angiosperm) forests in the central mountains of
grooved, glabrous, terminating in globose buds Taiwan. These forests are dominated by the families
ca. 4 mm diam. with imbricate, triangular scales. Fagaceae and Lauraceae, with Theaceae and Mag­
Leaves lanceolate to linear-lanceolate, (3–)5–10(–12) noliaceae also prominent. Characteristic genera of
cm long, 8–15 mm wide, straight or slightly curved trees are Cyclobalanopsis (= Quercus), Lithocarpus,
towards apex, tapering gradually to a petiolate and Machilus (= Persea), Cryptocarya, Schima, and Mag­
slightly twisted base ca. 5 mm long and an obtuse or nolia. The altitudinal range of this species rich ev-
sometimes acute apex; margins entire and smooth; ergreen forest type is from 700 to 1800 m a.s.l. and
midvein acutely raised adaxially, slightly raised or it exists under a markedly monsoonal subtropical
nearly flat abaxially; leaf colour mid green above climate.
(adaxially), pale green below; flushing leaves red-
dish pink or sometimes light green. Stomata on
Conservation
abaxial side, very small, in numerous irregular lines
on either side of midrib. Pollen cones axillary, ses- This species although not specifically targeted by
sile, solitary or in clusters of 2–3, becoming long loggers as it is generally of only small to moderate
cylindrical and lax at anthesis, 20–40 mm long, size, has suffered greatly from general deforestation
2–3 mm wide; microsporophylls arranged in steep in the period after World War II. Much of the pri-
spirals, triangular-apiculate, spreading, yellowish mary forest where P. nakaii occurred has gone, or it
green, with two sublateral, globular, yellow pollen has been turned into secondary vegetation unsuit-
sacs. Seed cones axillary, solitary on 2–10 mm long able for this species. A locality in Chianghua where
peduncles; receptacles subtended by (1–)2 small, W. R. Price collected the species in 1912 probably no
longer has it growing there; Flora of Taiwan, ed. 2, 1: 18 mm wide, acuminate; leaves more exposed to light
565 (1994) only mentions Nantou for its distribution. mostly linear-lanceolate or lanceolate, (4–)8–12(–15)
IUCN: EN (A1a-d, C2a) cm long, 6–15 mm wide, straight or slightly curved,
petiolate at a more gradually narrowing base, tapering
to an acute apex. Midrib acutely (sometimes obtusely)
Uses
raised adaxially, flat or obtusely (sometimes acutely)
No uses have been recorded for this species; its wood raised abaxially, usually narrower on adaxial side;
was logged with other trees, and if of good size and leaf colour dark green above, pale green below; new
shape would have been used for construction, car- leaves flushing red. Stomata very small, in numerous
pentry, etc. It is probably in cultivation, but it may irregular lines on abaxial (under) side on either side
masquerade under the name Podocarpus macro­ of midrib. Pollen cones axillary, solitary or in clusters 867
phyllus. of 2–3, sessile with several basal, carinate bud scales,
cylindrical, elongating to 25–40(–50) mm, 2–3.5 mm
wide; microsporophylls spirally arranged, triangular,
Podocarpus neriifolius D. Don, in Lambert, Descr. spreading, with two globose pollen sacs. Seed cones
Pinus 2: [21]. 1824. axillary, solitary on a 5–20 mm long peduncle; recep-
tacles with 2 subulate bracts (‘foliola’) 2–6 mm long
at base, swelling to become ellipsoid with a truncate
Etymology
distal end, 8–10 mm long, yellow turning orange-red
The species epithet means ‘leaf like that of the genus or bright red and succulent when ripe. Seeds solitary
Nerium’ [oleander]. at truncate end of receptacle, enclosed in a smooth
epimatium, green turning purplish green when ripe,
ovoid-oblong or ovoid-globose, (8–)10–15 × 7–8 mm.
Vernacular names
Seed proper ovoid, 6–10 mm long, slightly flattened.
Brown pine; podo bukit (Malay); numerous local
names are in use in Malesia (see De Laubenfels,
Taxonomic notes
Flora Malesiana, ser. 1, 10 (3): 401. 1988); bai ri qing
(Chinese); thông tre lá dài (Vietnamese). Pilger In Flora of China 4: 83 (1999) and in the World
(1903) cites kimerah and kiputri as names for the In- Checklist and Bibliography of Conifers (Farjon, 1998,
donesian island of Jawa, but these seem to apply to [2001]) P. annamiensis, described by N. E. Gray (op.
several species of ‘podocarps’ now in different gen- cit.) from Indochina, Myanmar [Burma] and Hainan
era. In Vanuatu the name ‘nuhuoto’ was recorded; in Island, is recognized as a distinct species, in China
Fiji (Viti Levu) ‘asimbolo’, (Vanua Levu) ‘kuasi’. restricted to Hainan. However, the stated differ-
ences with P. neriifolius are very small and seem to
be inconsistent. It appears that these differentiating
Description
character states (e.g. apiculate leaf apex, conspicu-
Shrubs or trees to 45 m tall, to 75(–100) cm d.b.h., ously raised midvein on both sides of the leaf in
bole straight, in large trees slightly fluted or rarely P. ­annamiensis) can be found in P. neriifolius from
buttressed at base. Bark smooth, thin, on large boles parts of its range beyond the areas where P. annami­
with narrow, longitudinal flakes or strips, light ensis is supposed to occur. Podocarpus annamiensis is
brown weathering grey; inner bark pinkish or red- therefore here treated as synonymous, in agreement
dish brown, fibrous. Branches spreading, forming a with recent treatments of the two taxa especially
rounded or domed crown. Foliage branchlets terete, concerning the conifers of Vietnam (Nguyen Tien
more or less grooved, glabrous, terminating in 2–5 Hiep & Vidal, 1996; Nguyen Tien Hiep et al., 2004).
(–8) mm long buds with spreading, triangular to lan- See also the Taxonomic notes under P. subtropicalis
ceolate outer scales. Leaves on juvenile plants short for an opposite view on which taxa to exclude from
petiolate, linear-lanceolate, 15–20(–25) cm long, or include with the widespread and variable species
17–26 mm wide, often curved and with undulating P. neriifolius. Gray (1958) described P. neriifolius var.
margins, abrubtly narrowed at base, long acuminate degeneri from Fiji (Viti Levu) as a distinct variety.
or acute. Leaves of mature trees usually much shorter Silba (1984) elevated it to a species, while De Lauben-
and narrower, shade leaves (6–)8–15 cm long, (8–)12– fels (1985) separated the two species into different
 sections, apparently based on slightly different sizes Dacrydium spp. and Podocarpus spp., and occasion-
of the ‘foliola’ or basal bracts on the receptacles. Such ally with Agathis spp. and/or Phyllocladus trichoma­
minutiae hardly justify classifications at the species noides, especially on mountain ridges, and with
level, considering variations, and are not evident in numerous angiosperm trees of the families Faga­
the specimens seen at Kew (K). In a revision of Fijiian ceae, Myrtaceae, Sapindaceae, Guttiferae (Clusiace­
conifers, Doyle (1998) has treated P. nerii­folius var. ae), Cunoniaceae, Rutaceae, and many others.
degeneri as a synonym of P. neriifolius. The narrower
leaves appear to justify varietal distinction and I fol-
Conservation
low Netta Gray in this case. Yet another new spe-
cies, P. epiphyticus, was described from Myanmar This species, while logged extensively as a valuable
868 [Burma] by De Laubenfels and Silba (1988). In the timber tree, is too widespread and common in most
protologue it is said to be “an epiphytic shrub” which areas to be classified as threatened with extinction
was taken from the type specimen label. The isotype under the IUCN criteria. Locally, its status may
specimen (J. Keenan 3081 at K) is a branchlet with be more severe depending on rates of logging and
typically broad juvenile foliage leaves belonging to levels of general deforestation. The species is listed
P. neriifolius. It was probably a young plant germi- on CITES Appendix III (at the request of Nepal in
nated and growing upon a tree, as conifers (and other 1975), which means that certificates or permits are
trees) sometimes do, but it can be categorically stated required to export its timber from that country.
that there are no obligate epiphytes among conifers. IUCN: status listed under varieties.

Distribution Uses
From E Nepal and Bangladesh through Myanmar This widespread species is an important timber tree
[Burma] and Indochina (including the Andaman Is- in several countries where it is common. Its excel-
lands), S and SW China, Malesia, Papuasia and Fiji. lent wood is used for house construction and car-
TDWG codes: 36 CHC-CQ CHC-GZ CHC-SC CHC- pentry and for making oars, spars, and masts of
YN CHH CHS-FJ CHS-GD CHS-GX CHS-HN CHS-JX sailing vessels. More specialized uses requiring high
CHS-ZJ CHT 40 ASS-ME ASS-MI BAN NEP 41 AND- grade timber are veneer, furniture making, cabi-
AN CBD LAO MYA THA VIE 42 BOR-BR BOR-KA net making, interior trim, household utensils, and
BOR-SB BOR-SR JAW LSI-BA LSI-ET LSI-LS MLY-PM wood carving. In New Guinea, Highland tribes use
MOL PHI SUL SUM 43 BIS NWG-IJ NWG-PN SOL-NO the bark to insulate walls of round houses. As far as
SOL-SO 60 FIJ known it is not in cultivation; in China and several
countries where plants under this name are intro-
duced and/or cultivated, these commonly belong
Ecology
to P. subtropicalis originating from Emei Shan [Mt.
Podocarpus neriifolius is the most widespread spe- Omei] in Sichuan, China, or perhaps to yet other
cies in the genus. It occurs as a scattered tree in ever­ similar species. This identity may also apply to a cul-
green primary broad-leaved forests, either in the tivar with variegated leaves that is occasionally seen
understorey or more commonly (and eventually) as labeled under P. neriifolius.
a canopy tree. Its altitudinal range in China is from
100 m to 1000 m a.s.l., but in tropical Malesia it oc- 2 varieties are recognized:
curs from near sea level to 2100 m a.s.l. It is usually
confined to well drained sites away from rivers, but Podocarpus neriifolius D. Don var. neriifolius.
occasionally it occurs in swampy forest. The soils Margbensonia neriifolia (D. Don) A. V. Bobrov
vary from nutrient-poor sands (‘kerangas’ in Bor- & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
neo), sandy clay or sandstone, to andesitic laterites Otd. Biol. 103 (1): 60. 1998. Type: Nepal [“Legi ad
(latosols) and ultrabasic soils over serpentine in New Sankoo,…Napaliae 1821”], N. T. Wallich 6052A
Guinea. In China and Indochina it is a ­component (lectotype BM, isolectotype K-W). Fig. 278, 279
in Lithocarpus-Castanopsis mixed evergreen for-
est. In Malesia this conifer is commonly associated Podocarpus neriifolius D. Don var. polyanthus Was­
with other podocarps e.g. Dacrycarpus imbricatus, scher, Blumea 4 (3): 455. 1941; Podocarpus polyanthus
(Wasscher) Gaussen, Trav. Lab. Forest. Toulouse Podocarpus neriifolius D. Don var. degeneri
T. 2, 1 (2, 21): 191. 1976 (nom. inval., Art. 33.2). N. E. Gray, J. Arnold Arbor. 39: 467. 1958.
Podocarpus decipiens N. E. Gray, J. Arnold Arbor. 36: Podocarpus degeneri (N. E. Gray) de Laub. ex
204. 1955; Podocarpus neriifolius D. Don var. decipi­ Silba, Phytologia Mem. 7: 61. 1984; Margbensonia
ens (N. E. Gray) Silba, J. Int. Conifer Preserv. Soc. 7 degeneri (N. E. Gray) A. V. Bobrov & Melikyan,
(1): 35. 2000. Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1):
Podocarpus annamiensis N. E. Gray, J. Arnold Arbor. 60. 1998. Type: Fiji: Viti Levu, Mba, Nandarivatu,
39: 451. 1958. O. Degener DA 14272 (holotype A).
Podocarpus annamiensis N. E. Gray var. hainanensis
Gaussen, Trav. Lab. Forest. Toulouse T. 2, 1 (2, 21):
Description
175, 210. 1976 (nom. inval., Art. 37.1). 869
Podocarpus epiphyticus de Laub. & Silba, Phytologia Shrubs or small trees. Terminal buds large, to 8 mm
64: 290. 1988. long, ovoid, outer scales acuminate to long attenuate
Podocarpus neriifolius D. Don var. penibukanensis or apiculate. Leaves on mature trees 6–12 cm long,
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 34. 2000. 6–10 mm wide, linear, acute. Leaves on juvenile
Podocarpus neriifolius D. Don var. staintonii Silba, plants to 18 cm long and 17 mm wide.
J. Int. Conifer Preserv. Soc. 7 (1): 36. 2000.
Distribution
Description
Fiji: Vanua Levu, Viti Levu.
Trees to 45 m tall. Terminal buds small, 2–4 mm TDWG codes: 60 FIJ
long, with spreading, triangular to lanceolate outer
scales. Leaves on mature trees 8–15 cm long, 12–18
Ecology
mm wide, acuminate; leaves more exposed to light
mostly linear-lanceolate or lanceolate, (4–)8–12(–15) Podocarpus neriifolius var. degeneri occurs in ever-
cm long, 6–15 mm wide. Leaves on juvenile plants green forests, at altitudes between 40 m and 1050 m
15–20(–25) cm long, 18–26 mm wide a.s.l. In one locality it was found growing with Musa,
Casuarina, Myrtaceae, etc. in dense forest; other
records mention “dry forest, along stream”, so it is
Distribution
dependent on a permanent supply of water. Where
From E Nepal and Bangladesh through Myanmar precipitation increases with altitude, this podocarp
[Burma] and Indochina (including the Andaman Is- will be mixed with other rainforest trees away from
lands), S and SW China, Malesia, Papuasia and Fiji. streams.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-SC CHC-
YN CHH CHS-FJ CHS-GD CHS-GX CHS-HN CHS-JX
Conservation
CHS-ZJ CHT 40 ASS-ME ASS-MI BAN NEP 41 AND-
AN CBD LAO MYA THA VIE 42 BOR-BR BOR-KA Podocarpus neriifolius was assessed as “LRlc” (Least
BOR-SB BOR-SR JAW LSI-BA LSI-ET LSI-LS MLY- Concern) for Fiji by M. F. Doyle in Farjon & Page
PM MOL PHI SUL SUM 43 NWG-IJ NWG-PN SOL-NO (1999) and this pertains to both varieties.
SOL-SO 60 FIJ IUCN: LC

Conservation
Podocarpus nivalis Hook., Icon. Pl., n.s., 2: t. 582.
IUCN: LC 1843. Type: New Zealand: [locality unknown],
W. Colenso s.n. (holotype K). Fig. 280, 281

Etymology
The species epithet means ‘growing in or near snow’
and refers to the alpine habitat of this species.
 (Chionochloa rubra) it is often associated with Hebe,
Vernacular names
Olearia, Dracophyllum, Gleichenia, and the moss
Mountain totara, Snow totara Racomitrium, etc. In scrubland it can be found with
the conifers Halocarpus bidwillii and Phyllocladus
trichomanoides var. alpinus, often interspersed with
Description
ericoid dwarf shrubs and scattered grasses in a
Decumbent, sprawling or more or less erect shrubs, lichen-­rich ground cover.
20–40 cm to 2 m tall, multi-stemmed. Foliage
branches spreading or ascending, in decumbent
Conservation
shrubs with creeping whip shoots, forming mat-
870 ted shrubs of a few square meters in extent. Termi- IUCN: LC
nal buds very small, with imbricate scales, hidden
among leaves. Leaves crowded, spirally arranged,
Uses
spreading at narrow to wide angles from shoot, 3–10
mm long, 2–2.5 mm wide, boat-shaped (naviculate) Mountain totara is fairly common in cultivation, es-
to ovate-linear, curved outwards or down on some pecially for rock gardens where prostrate forms will
branchlets, narrowly decurrent at base, obtuse or grow readily over rocks, banks and low walls. The
mucronate at apex, thick coriaceous, rigid, canal- more upright forms are suitable for hedges lower
iculate (grooved) adaxially (above), with an obtuse than 1 m and take regular clipping well. This shrub
midrib abaxially. Stomata on abaxial (lower) side of is quite hardy depending on provenance and grows
leaf, on either side of midrib below, in intermittent on almost any soil. No cultivars are known of this
white lines. Pollen cones axillary, on short pedun- species and plants in the trade are propagated from
cles with several small, obtuse bracts, solitary or cuttings; as the species is dioecious these usually do
with 2–3 or more together, very slender cylindical, not produce receptacles (‘berries’).
10–15 mm long, 1–1.5 mm wide at anthesis; micro-
sporophylls peltate, apiculate, very small, with two
relatively large, globose pollen sacs. Seed cones axil- Podocarpus novae-caledoniae Vieill., Ann. Sci.
lary, nearly sessile, solitary; receptacles originating Nat. Bot., sér. 4, 16: 56. 1862. Type: New Caledonia:
from a short axis with two fused bracts, swelling to Grande Terre, Province Sud, Païta, E. Vieillard
sub-globose, 8–10 × 8 mm, becoming succulent and 1266(b) (holotype P). Fig. 282, 283
scarlet. Seeds including the epimatium ovoid with
a constricted distal end, 4–5 × 3 mm, lustrous light Podocarpus beecherae de Laub., New Zealand J. Bot.
green. Seed proper not observed. 41: 715. 2003.

Distribution Etymology
New Zealand: North Island, South Island, Stewart The species epithet refers to New Caledonia, where
Island. the species is endemic.
TDWG codes: 51 NZN NZS
Vernacular names
Ecology
No common names have been recorded of this spe-
Podocarpus nivalis occurs at higher altitudes in the cies.
sub-alpine to alpine scrubland and tussock grass-
land of the New Zealand mountains. Its altitudinal
Description
range varies from 1000 m to 2500 m a.s.l. depend-
ing on local climate and exposure. It is a species Shrubs 1–2 m tall, rarely a small tree to 6 m tall; dense-
that commonly creeps low over rocks, but when ly branched, erect and rounded or more elongated
competing with tall grasses or in tall scrubland it in small trees. Bark fibrous, light brown weathering
may grow to a more erect bush, sometimes several grey. Foliage branchlets numerous, slender, terete,
meters wider than tall. In open tussock grassland glabrous with minute grooves between leaf bases,
terminating in small, ovoid or subglobose buds of these stated differences in herbarium specimens
with imbricate, appressed, triangular, weakly keeled, at K has shown that these distinctions either do not
1–2 mm long bud scales. Leaves on juvenile plants hold or are at best unconfirmed (colour of recepta-
longer than on mature plants, up to 12 cm. Leaves on cle). A collection not known to De Laubenfels from
mature plants (2.5–)4–8(–10) cm long, 3–5(–6) mm the Île des Pins (Pic N’Ga) was found away from any
wide, linear, straight or slightly falcate, gradually or stream in maquis and had red receptacles. The yel-
sometimes more abruptly widening from a petiolate lowish green receptacles found at the type locality of
base, mostly with two parallel sides up to a gradually P. beecherae could well be a fully grown but yet unripe
tapering, acute or obtuse apex. [Shortest leaves may stage [when I visited the site in November 2005 the
differ in being either narrowly lanceolate and acute small receptacles were glaucous but immature]; this
or sometimes slightly wider above the middle and is seen in several other species in the genus, where 871
then obtuse.] Midrib on adaxial (upper) side often the colour turns from yellowish green to yellow, or-
running in a shallow V-shaped depression, form- ange, red and even purple eventually. Receptacle col-
ing a groove or a faint raised rib only proximally, on our is not a sufficient character to base a species on.
abaxial side obtusely raised; leaf texture coriaceous; Besides, a mature greenish yellow receptacle would
leaf colour lustrous light green above, dull light green be unlikely in the genus; its colour is an adaptation
below. Stomata numerous, small, in two wide bands to bird dispersal and therefore has to be quite dis-
of intermittent lines on abaxial side. Pollen cones ax- tinct from the foliage when the seed is ripe to be
illary, sessile, solitary or 2–3 together, narrowly cy- effective. The leaf shapes of the type of P. beecherae
lindrical, 7–18 mm long, 2–2.5 mm wide at anthesis, and other collections cited by De Laubenfels fall en-
yellow turning brown; microsporophylls very small, tirely within the variation seen in those specimens
triangular, with two sublateral, globose pollen sacs. of P. novae-­caledoniae he did not include in his new
Seed cones axillary on slender peduncles 5–15 mm species. Podocarpus beecherae is therefore consid-
long, solitary; receptacles subtended by two small, ered a synonym of P. novae-caledoniae.
recurved bracts (‘foliola’) and consisting of an axis
with 2–3 fertile bracts which amalgamate and be-
Distribution
come swollen and succulent, 9–12 mm long, 6–8
mm wide, yellowish green to orange or (dark) red. New Caledonia: Grande Terre (Province Sud), Île
[In some populations receptacles seem to stay green- des Pins
ish but it is not clear if they do not become more TDWG codes: 60 NWC
coloured eventually.] Seeds solitary or two on a re-
ceptacle, including the epimatium irregularly ovoid,
Ecology
6–8 × 5–6 mm, with or without a small crest, whitish
green or pruinose, ripening to light brown, smooth Podocarpus novae-caledoniae is restricted to the
and lustrous. Seed proper not observed. ultramafic rock formations of the southern part of
New Caledonia. It occurs often along streams and
small rivers, but not in regular contact with their
Taxonomic notes
waters, and is also but less frequently found on
De Laubenfels (op. cit.) described a new species, rocky slopes away from streams, at altitudes from
Podo­carpus beecherae, from a locality at the Col de near sea level to ca. 700 m a.s.l. It is associated with
Yaté (Djuru Déré, near a Catholic shrine marked by shrubs and dwarfed trees in the ‘maquis minier’ veg-
a large cross) growing in ‘maquis minier’ on eroded etation, where it normally stays a low, dense shrub
serpentine. It has according to the protologue “light often forming bushy thickets. Among trees like Aga­
greenish yellow and glaucous” receptacles, distinct this ovata, and to a lesser extent Dacrydium arau­
from the red receptacles of P. novae-caledoniae. The carioides and Casuarina sp., it may be forced up to
leaf shapes of the two species are also said to differ, a small tree 4–6 m tall, with a narrow crown. Rain-
with more lanceolate leaves in P. beecherae and more fall is high in most areas, especially on SW exposed
linear shapes in P. novae-caledoniae. They are claimed mountain slopes, but the vegetation remains mostly
to occur in different habitats, with P. novae-caledo­ open due to extreme soil conditions not suitable for
niae termed a “semi-rheophyte” while P. beecherae high plant density, giving this undemanding conifer
occurs in maquis on dry rocky sites. Closer scrutiny an opportinity to thrive.
 on adaxial (upper) side, obtusely raised or nearly
Conservation
flat on abaxial side. Stomata very small, in intermit-
IUCN: LC tent lines forming two white bands separated by a
green midrib on abaxial face. Pollen cones axillary
towards ends of leafy shoots, in groups of 2–4 per
Uses
leaf, sessile with small perular scales at their base,
No uses have been recorded of this species; it should cylindrical, 10–20(–25) mm long, 2.5–3.5 mm wide,
be anemable to cultivation for gardens in warm cli- cream coloured; microsporophylls deltoid, with
mate regions, requiring very little in the way of soil an acute, erose-denticulate apex, each bearing two
fertility. large, globose pollen sacs. Seed cones axillary, soli-
872 tary on short peduncles; receptacles formed of an
axis with two unequal bracts, fused and swelling to
Podocarpus nubigenus Lindl., J. Hort. Soc. London 7–8 × 6–8 mm, red ripening to dark purple, succu-
6: 264. 1851. Type: Chile: Los Lagos, Isla de Chiloé, lent. Seeds including the epimatium ovoid, 8–9 ×
W. Lobb 80 (holotype not located, isotype K). Fig. 6–7 mm, not crested, dark purplish green ripening
284, 285 to purple. Seed proper not observed.

Etymology Distribution
The species epithet is a compound adjective from S Argentina: Neuquén, Santa Cruz; S Chile: Aisén,
Latin nubilus = ‘dark, cloudy’ and genus = ‘born’ or Los Lagos, Magellanes.
‘produced’, and refers to the dark purple seed (and TDWG codes: 85 AGS-NE AGS-SC CLS-AI CLS-LL
perhaps receptacle). CLS-MG

Vernacular names Ecology

Manío, Manío macho, Manío hembra, Pino amarillo Podocarpus nubigenus is a component of mixed tem-
(Chile) perate rainforest, occurring from sea level to 1000 m
a.s.l. It may grow in a forest type dominated by coni-
fers, e.g. Fitzroya cupressoides forest with P. nubi­
Description
genus, Saxegothaea conspicua and Pilgerodendron
Trees to 25 m tall or more; trunk to 2 m d.b.h., mo- uviferum, or in forest dominated by angiosperms,
nopodial, erect. Bark smooth in young trees, on in particular Nothofagus nitida. Other angiosperm
large trunks shallowly fissured, scaly, exfoliating trees with which it frequently occurs are Drymis
in elongated flakes, cinnamon or purplish brown, winteri, Laureliopsis philippiana, Nothofagus betulo­
weathering grey. Branches in younger trees ascend- ides, and various species in the family Myrtaceae.
ing, forming a pyramidal crown, in old trees fewer ­Podocarpus nubigenus is often found in water-logged
heavy branches form a more open, irregular crown. soil or near streams on acidic substrates, particularly
Foliage branches stout or slender, terete, grooved in the coastal mountains and on the islands in the
between leaf bases. Terminal buds small, up to southern archipelago. It may also occur in montane
6 mm long on vigorous shoots, with erect or slightly shrub vegetation including Embothrium coccineum,
spreading, free, lanceolate scales with acute apices. Ovidia andina, and Desfontainia spinosa, on a sub-
Leaves on saplings and vigorous shoots slightly larg- strate of volcanic ash in the Andes.
er than on lateral twigs of mature trees. Leaves on
mature trees patent, 1.5–3 cm long, linear-lanceolate,
Conservation
straight or very slightly falcate, 2–4 mm wide, more
or less abruptly narrowing to a decurrent base, rigid- Although widespread, this species is still at risk from
ly coriaceous, gradually tapering to a pungent apex, logging, both for firewood and its high quality tim-
dark green with a lighter midrib above, two white ber; while young trees are often cut as Christmas
bands below. Midrib raised but fading ­towards apex trees. Changes in land use, especially conversion of
natural forest to plantations of Eucalyptus and Ra-
Vernacular names
diata pine, further reduce its extent of occurrence
(EOO) and area of occupancy (AOO). Due to its Pino de pasto (Colombia); Pinete (Venezuela)
wide distribution and large subpopulations still in
existence, Podocarpus nubigenus does not at present
Description
meet the criteria for a threatened species.
IUCN: NT Trees to 20–25(–30) m tall, to 1.5 m d.b.h., mono­
podial, erect, branching densely, at high altitudes
a stunted bush or tree. Bark thin, eventually scaly,
Uses
yellowish brown weathering light grey. Foliage
This species is a valuable timber tree and is being branches terete, with prominent leaf scars and de- 873
logged for its timber of larger size, but also cut for current grooves. Terminal vegetative buds initially
firewood by the rural population. Young trees are sub­globose, 3–5 mm wide with imbricate, short,
cut from the forest to be used as Christmas trees. It rounded or ovate scales 3–4 mm long which often
is in cultivation in the milder Atlantic parts of Eu- spread out later; axillary buds globose, with imbri-
rope, especially in large gardens with tree collections cate, thick, rounded, carinate scales 2–3 mm long.
in Ireland and southern England. It should be used Leaves on saplings often much larger than on ma-
more in horticulture, also in view of potential threat ture trees, to 15 cm long and 20 mm wide, linear-­
to the species in the wild; instructions for propaga- lanceolate, often slightly falcate. Leaves on mature
tion can be found in Gardner et al. (2006). trees lanceolate to linear-lanceolate, straight (some-
times slightly curved), petiolate at base, gradually
narrowing towards base as well as the acute or more
Podocarpus oleifolius D. Don, in Lambert, Descr. or less obtuse apex, extremely variable in size de-
Pinus 2: [20]. 1824. Type: Peru: [“in Peruvia ad pendent on growth of shoot, usually 4–9 cm long
Pillao et Panao”], R. Ruiz & J. A. Pavón y Jiménez and 7–12 mm wide, but as small as 1.5 cm long and
s.n. (lectotype BM, designated here). 5 mm wide on some trees. Midrib on adaxial (up-
per) side not elevated above surface of lamina but
Podocarpus macrostachyus Parl., in Candolle, Prodr. forming a narrow medial rib in a groove, on abax-
16 (2): 510. 1868; Podocarpus oleifolius D. Don var. ial side a more or less prominent ridge, often with a
macrostachyus (Parl.) J. T. Buchholz & N. E. Gray, central groove continuing to apex. Pollen cones axil-
J. Arnold Arbor. 29: 140. 1948. lary, solitary, sessile, expanding to 25–30 mm long,
Podocarpus oleifolius D. Don var. costaricensis 3–4 mm wide; microsporophylls imbricate, with
J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 140. rounded or apiculate apex, bearing two pollen sacs
1948. near the base. Seed cones axillary, on short pedun-
Podocarpus oleifolius D. Don var. trujillensis cles; receptacles 6–10 mm long, swelling to become
J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 141. succulent and red. Seeds including the epimatium
1948. ovoid-globose, 7–8 mm long, with an inconspicuous
Podocarpus oleifolius D. Don var. equadorensis Silba, crest, olive green. Seed proper not observed.
Phytologia 68: 69. 1990.
Podocarpus monteverdeensis de Laub., Brenesia 33:
Taxonomic notes
120. 1991.
Podocarpus ingensis de Laub., Bol. Lima 73: 58. 1991. This species was mentioned by D. Don in Lam-
Podocarpus ballivianensis Silba, J. Int. Conifer Pre- bert (1824) to be found in Chile based on a collec-
serv. Soc. 15 (1): 1. 2008. tion by R. Ruiz & J. A. Pavon y Jiménez, but it is not
known from further south than Peru and Bolivia.
The original material is in BM and was collected by
Etymology
Ruiz & Pavon in Peru at “Pillas”; of two sheets the
The species epithet oleifolius compares the leaves one labeled “in Peruvia ad Pillas et Panao” is here
with those of olives (Olea L.). designated the lectotype. Buchholz & Gray (op. cit.)
described P. oleifolius var. costaricensis from Vol-
can Poas; the type (H. Pittier 822) as well as other
 c­ ollections from there have short leaves and small, pensis and Q. cortezii, also with Magnolia sp., Per­
crested seeds. Similar specimens with small leaves sea sp., Cornus diciflora, Ilex sp., Cyatheaceae, and
are known from other localities in Central Amer- many bromeliad epiphytes and an epiphytic cactus.
ica and it is doubtful that they all belong to a sin- At the highest mountain ridges P. oleifolius occurs
gle taxon distinct from the more common form of in an evergreen elfin forest with a canopy only 2–8
the species. They are known from both high and m tall and here it is also a stunted bush or tree. In
middle elevations. The relatively recently described Venezuela it has also been found on the tepuis, high
new species P. monteverdeensis de Laub., from the sandstone table mountains with on their summit
reserve Monteverde in Costa Rica and found in a plateaus a mozaic vegetation of grasses, herbs and
“mountain top swamp forest” appears to be merely ferns, and dwarf forest or scrub.
874 a form with narrow leaves with a more or less at-
tenuate apex. This taxon and P. macrostachyus Parl.
Conservation
have the same overall leaf shape variation and very
similar midvein morphology as well as vegetative IUCN: LC
buds, and with no other characters to differentiate
them, they are here treated as synonyms of P. olei­
Uses
folius. Another species, P. ingensis, was described by
De Laubenfels (op. cit.) from Bolivia and Peru, said Only larger trees of this species growing in montane
to differ from P. olei­folius in foliage buds with more rainforest are being logged for timber, which is used
or less free scales and with lanceolate leaves widest in construction of houses and in furniture making.
below the middle. The pollen cones were unknown This species is very rare in cultivation and restricted
and the seed cones only seen as immature. Despite to a few botanic gardens.
several more recent collections made by A. Fuentes
and others (MO) under this name, this situation has
not improved. This species is doubtfully distinct and Podocarpus palawanensis de Laub. & Silba,
its described characters likewise fall within the mor- Phytologia 64: 291. 1988. Type: Philippines:
phological variation of P. oleifolius that can be ob- Palawan, Pagdanan Range, Ibangley [“Brookside
served in much of its wide geographical range. Hill”], C. E. Ridsdale SMHI 1502 (holotype L).

Distribution Etymology
S Mexico; Central America; Bolivia; Colombia; The species epithet refers to the island of Palawan,
Equador; Peru; Venezuela. Philippines.
TDWG codes: 79 MXC-PU MXG-VC MXS-GR MXS-
MI MXS-OA MXT-CI 80 COS ELS GUA HON PAN 82
Vernacular names
VEN 83 BOL CLM ECU PER
No common name has been recorded for this spe-
cies.
Ecology
Podocarpus oleifolius is a montane to high mon-
Description
tane species occurring in primary semi-deciduous
and evergreen forests. Its altitudinal range is (825–) Trees to 7 m tall. Buds on vegetative branchlets 4–6
1200–3300 m a.s.l. and it is most frequent above mm long and 4–5 mm wide at their base, with tri-
2000 m a.s.l. In some high valleys and on mountain angular, erect or slightly spreading outer scales.
summits at moderate elevation it grows in swamp Leaves linear-lanceolate, 10.5–18.5 cm long, 8–11
forest. In tall montane forests it is associated with mm wide, narrowing to a 5–7 mm long petiolate
e.g. Cedrela, Persea, Quercus, Podocarpus, Magnolia, base and an acute apex. Midrib on adaxial (upper)
and Symplocus and various species in Lauraceae. At side acutely raised. Pollen cones axillary, sessile, sol-
a higher altitude in Central America it was found itary, cylindrical, 35–45 mm long, 6.5–8 mm wide;
in cloud forest with the dominants Quercus xala­
­ icrosporophylls with a 4 mm long, lanceolate apex
m Podocarpus pallidus N. E. Gray, Bishop Mus. Bull.
and two basal pollen sacs. Seed cones not observed. 220: 46. 1959. Type: Tonga: Tongatapu Group,
‘Eua, ‘Ohonua, [“western side of plateau region”],
[This brief description has been translated from the H. E. Parks 16212 (lectotype F).
Latin description given in the protologue; no other
information being available.]
Etymology
The species epithet means ‘pale’ and probably refers
Taxonomic notes
to the pale green leaves.
This species was first described and named in 1988,
based on a single collection, C. E. Ridsdale SMHI 1502 875
Vernacular names
(holotype L, isotype K) with aberrant pollen cones.
This is unsatisfactory and a critical comparison with uhiuhi (Tongan name)
other species in the region is needed, but hampered
by the lack of collections particularly from Palawan.
Description
It is a species that has here been given the benefit
of the doubt, but may be synonymous with P. poly­ Shrubs or small trees to 10 m tall, usually monopo-
stachyus. dial or branching low; trunk to 25 cm d.b.h. Bark
light brown weathering grey; inner bark reddish
brown. Foliage branchlets ascending or spread-
Distribution
ing, terete, with fine grooves or ridges, terminat-
Philippines, Palawan (Pagdanan Range). ing in small ovoid-conical buds with 2–6 mm long,
TDWG codes: 42 PHI spreading or recurving and acuminate outer scales.
Leaves on juvenile plants 10–15 cm long, 12–17 mm
wide. Leaves on mature plants linear-lanceolate
Ecology
to linear, (2.5–)4–12 cm long (very variable on the
Closed tropical evergreen rain forest dominated by same branch), 5–11 mm wide, straight or slightly
angiosperms. falcate, gradually tapering to a petiolate base and to
an acute or acuminate, sometimes mucronate apex;
margins occasionally slightly revolute; leaf colour
Conservation
lustrous light green above, pale green below. Midrib
Not known from any other than the type loca- on adaxial (upper) side narrow (0.3–0.4 mm wide),
tion, which is called Ibangley Brookside Hill in the acute but sometimes fading towards leaf apex, on
Pagdanan Range at only 40 m altitude. A map in abaxial side wider (to ca. 1 mm) and obtuse or flat-
the Philippines Red Data Book showing the extent tened. Stomata small, in numerous intermittent
of natural forest cover on the island of Palawan at lines on abaxial side on either side of distinct mid-
present demonstrates an extreme rate of deforesta- rib. Pollen cones axillary, solitary, sessile, subtended
tion, much of which has occurred in recent decades. by triangular bud scales, long cylindrical, 25–40 mm
It is not known if this has affected the type location, long, 3–4 mm wide at anthesis, pale yellowish white;
and thereby this species, but assuming that it was microsporophylls triangular, acute, with scarious
present in places outside the type location, it is likely margins and two sub-basal pollen sacs. Seed cones
to have suffered serious decline. This species is not axillary, solitary on 5–15 mm long, stout peduncles;
within a protected area. receptacles subtended by two ca. 2 mm long, curved
IUCN: CR [B1, B2 a-b (iii)] bracts (‘foliola’), and formed of an axis with 2–3
fused bracts of unequal size, swelling to a succulent,
red to blackish red body 10–13 mm long, ca. 10 mm
Uses
wide at distal end. Seeds single, including the epima-
No uses are specifically known of this species. Pre- tium ovoid to subglobose, 10–13 × 8–10 mm, with or
sumably it will have been used or is being used for without an obscure distal ridge, blue to brown with a
firewood locally. glaucous bloom. Seed proper not observed.
 fissured on larger stems, brown weathering grey.
Distribution
Branches often ascending from near the ground,
SW Pacific: Tonga (‘Eua and ‘Uta Vava’u). but in closed forest trees have a clear bole to some
TDWG codes: 60 TON height, forming a broad crown. Foliage branches
sub-verticillate, ascending to erect, slender, terete,
with fine grooves running down from slightly el-
Ecology
evated leaf bases, yellowish green turning light
Podocarpus pallidus occurs in tropical low forest, brown. Terminal buds broadly ovoid to subglobose,
dominated by angiosperms. This species is usually 4–6 mm long and wide, with numerous apiculate
scattered in forest patches along the plateau escarp- scales with minutely denticulate margins and free
876 ment and in ravines or gullies of two islands in the tips, giving buds a bristly appearance; axillary buds
Tonga Archipelago situated some 325 km apart. It smaller, globose. Leaves on juvenile plants or sap-
has been found on limestone cliffs on ‘Uta Vava’u lings larger than on mature plants, to 12 cm long and
and occurs from around 50 m to ca. 250 m a.s.l. 5 mm wide. Leaves on mature plants patent to more
or less erect, (2.5–)4–8(–9) cm long, 2–3 mm wide,
narrowly linear, straight or slightly falcate, with a
Conservation
short petiolate base; margins slightly revolute, very
This species appears to occur on two islands, ‘Eua gradually tapering to an acute-pungent apex, thin
and ‘Uta Vava’u, of the Tonga Archipelago. No col- coriaceous, green above, pale green below; flushing
lections are known from Tongatapu, Tofua or Late, leaves light green. Midrib on adaxial (upper) side of
other islands in the archipelago with low forested leaves a shallow groove often fading towards apex,
mountains and plateaux. Because this species is a on abaxial side a narrow ridge, continuous or raised
minor component in broad-leaved tropical forest, in lower part of leaves only. Stomata very small, in
it is assumed there are fewer than 500 mature trees numerous intermittent lines on abaxial side. Pollen
on each island. We have no information about pos- cones in clusters of (3–)4–6 or more due to corym-
sible decline. This species is not known from any bose branching, on axillary, slender, angular stalks
protected area. 5–15 mm long; individual cones axillary to a 2–3
IUCN: VU (D 1, 2) mm long, lanceolate, acute bract, sessile and sub-
tended by small, triangular, carinate-apiculate bud
Uses scales, cylindrical, 5–10 mm long, 1.5–2 mm wide;
microsporophylls triangular, with minutely den-
No uses have been recorded of this species.
ticulate margins and two globose pollen sacs. Seed
cones axillary, appearing with new leaves, solitary
Podocarpus parlatorei Pilg., in Engler, Pflanzenr. on 3–8 mm long peduncles; receptacles composed
IV.5 [18]: 86. 1903. Podocarpus angustifolius Parl., in of an axis with 2–3 fused, swollen bracts, 4–5 mm
Candolle, Prodr. 16 (2): 512. 1868, non Griseb. (1866. long and red when mature. Seeds globose, including
Type: Bolivia: [locality not stated], T. Bridges s.n. the epimatium 5–6 mm long, 4–5 mm wide, with or
(holotype not located, isotype K). without an inconspicuous crest.

Etymology Distribution
This species has been named after the Italian bota- Argentina (Jujuy, Salta, Tucumán, Catamarca); Bo-
nist Filippo Parlatore (1816–1877). livia (Cochabamba, La Paz); marginally in Peru
(­Sierra de Chaglla).
Vernacular names TDWG codes: 83 BOL PER 85 AGW-CA AGW-JU
AGW-SA AGW-TU
pino del cerro, pino blanco (Argentina, Bolivia)

Description Ecology
Shrubs or stunted trees to 12 m tall, occasionally to Podocarpus parlatorei occurs in montane scrubland
20 m tall; trunks of trees to 1.5 m d.b.h. Bark slightly and forest at altitudes between 1000 m a.s.l. and the
upper limit of trees between 2400 m and 3000 m simple furniture and tool handles. In Argentina it
a.s.l. This species may grow into a tree of maximal was heavily used for paper pulp in the 1970’s and
20 m height in riverine forest at lower altitudes, but 80’s; now (1990’s-2000’s) it is occasionally logged for
it is more often a shrub or stunted tree, branching local (domestic) uses only. In Bolivia there is a pro-
near the base. It is commonly associated in the for- gramme to exploit the species to use the wood for
est with Meliaceae, Myrtaceae (typically with Alnus local housing.
-Betulaceae- in NW Argentina and S Bolivia) and
other angiosperm trees; on cliff sides and in steep
rocky gullies it is often the dominant shrub and can Podocarpus pendulifolius J. T. Buchholz &
be quite abundant. In valleys it is locally abundant N. E. Gray, J. Arnold Arbor. 29: 138. 1948. Type:
on old river terraces or on river banks, often now in Venezuela: Cordillera do Mérida, Paramo de 877
forest relicts. It is a pioneer species with a key role la Negra, above La Grita, J. A. Steyermark 57115
in the forest dynamics. Its seeds or fruits are a food (holotype F). Pl. 38
resource for endangered and of restricted range bird
species, such as Penelope dabbenei (Cracidae) and
Etymology
Amazona tucumana (Psittacidae).
The species epithet refers to the drooping or pendu-
lous leaves.
Conservation
This species was listed as Data Deficient in the Glo-
Vernacular names
bal Red List of Conifers (Farjon & Page, 1999) and
since no reassessment has yet taken place, this is pino carbón, pino hayuco (Venezuela)
its status in the IUCN Red List at present. It has in
the past been considered threatened with extinc-
Description
tion and it was listed in 1975 on CITES Appendix
I, which prohibits export of any of its parts, includ- Trees to 20 m tall, often smaller (2 m tall trees can be
ing timber. In NW Argentina this species was heav- found fertile), d.b.h. to 50 cm. Bark thin, with hori-
ily used for paper pulp in the 1970’s and 80’s. Now zontal and vertical shallow fissures, reddish brown.
this used has been stopped. Trade in this species, if Branches spreading, forming a dense crown. Foli-
it exists, is unlikely to involve timber in any quan- age branchlets stout, spreading or ascending, terete,
tity that could be of commercial importance. This grooved between remote leaf bases, yellowish green
species rarely attains tree sizes over 15 m tall, but in turning light brown. Terminal buds subglobose, 3–5
Argentina trees from 15–20 m have been recorded. mm wide, often shorter than wide, with imbricate,
It is a species with high ability to recruit. For exam- broadly ovate, carinate scales with scarious margins.
ple, it is common in secondary forest along roads, Leaves strongly drooping or pendulous; leaves on
abandoned farms, and grazing areas where the use the upperside of the shoot curved down above a pet-
of fire was diminished or halted. Deforestation of a iolate base, 5–12 cm long, lanceolate-linear, 7–11 mm
general kind, in which this species, where it occurs wide, straight or slightly falcate, gradually tapering
as a tree, will be logged with the rest, could be of at both ends; margins nearly flat; apex acute; tex-
conservation concern. To change current status of ture thin coriaceous; leaf colour green on both sides.
the species for the IUCN Red List, it is required to Midrib obtusely raised and continuous to apex on
assess the species density and regeneration along its both sides of leaf. Stomata very small, in wavy, in-
distributional range, to evaluate the effect of differ- termittent lines on abaxial side in broad bands on
ent land practices on the species, and to determine either side of midrib. Pollen cones axillary, solitary,
the species habitat requirements to recruit. sessile, with obtuse perular scales at base, cylin-
IUCN: DD drical, 20–40 mm long, 6–7 mm wide at anthesis;
microsporophylls broadly ovate, obtuse with erose-
denticulate margin, bearing two globose pollen
Uses
sacs. Seed cones axillary, solitary on 7–10 mm long
The wood of this species is at present used locally peduncles; receptacles 8–9 mm long, swelling and
for firewood, fence posts and domestic articles, e.g. becoming red to dark purple, often pruinose. Seeds
 2
1

878

Pl ate 3 8 . Podocarpus pendulifolius; Podocarpus salicifolius. 1. Branch with leaves and seed cones.
2. Seed cone. 3. Habit of tree. 4. Branch with leaves and pollen cones. 5. Seed cone. 1–2 = P. pendulifolius; 3–5
= P. ­salicifolius.
single, attached at distal end of receptacle, including Podocarpus perrieri Gaussen & Woltz, Bull. Soc.
the epimatium 8–9 mm long, 6–7 mm wide, ovoid- Hist. Nat. Toulouse 111: 319. 1975. Podocarpus
globose, glaucous pruinose, with a small apiculate rostratus L. Laurent var. perrieri (Gaussen & Woltz)
crest or nearly obtuse. Seed proper not observed. Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39. 2000.
Type: Madagascar: Antananarivo Prov., Toamasina,
Forêt d’Andasibé, J. M. H. A. Perrier de la Bathie
Distribution
17109 (holotype P).
NW Venezuela: Cordillera do Merida.
TDWG codes: 82 VEN
Etymology
This species was named after Henri Perrier de la 879
Ecology
Bathie, who collected the type specimen.
Podocarpus pendulifolius occurs in montane mixed
low-canopy rainforest at elevations between 1400
Vernacular names
m and 3000 m a.s.l. At the highest elevations it is
part of a dwarfed forest on slopes immediately be- No common names have been recorded for this spe-
low the paramo vegetation, an alpine grassland rich cies.
in herbs. Young trees are said to be locally abundant
in this habitat. It is less common in the taller forests
Description
below these stunted forests, where it can occasion-
ally attain a height of 20 m. Trees to 30 m tall; trunk to 80 cm d.b.h. Bark thin,
becoming scaly on large trunks, light brown, weath-
ering grey. Branches ascending and spreading, in
Conservation
old trees forming an umbrella-shaped crown. Foli-
Based on distribution data from herbarium speci- age branchlets numerous, ascending, slender, terete,
mens, the extent of occurrence (EOO) is less than finely grooved between leaf bases, terminating in
5000 km2 and the area of occupancy (AOO) is less small buds with imbricate, obtuse outer scales and
than 500 km2; the species is only known from two acute or apiculate inner scales with free but incurved
populations ca. 100 km apart. Trees of some size are apices. Leaves on saplings and young trees similar
known to be valued and used for building houses, to those on mature trees, more remotely placed on
so there must be logging targeted at this species. No branchlets and at the larger end of the size range.
(sub)populations have been reported in protected Leaves on mature trees crowded, spreading to near-
areas. ly erect, 0.7–2(–2.5) cm long, rigid, linear, straight
IUCN: EN [B1, B2 a-b (iii)] or slightly curved, 1–2 mm wide, irregularly oval
in cross-section, narrowing slightly to a decurrent
base; apex pungent. Midrib absent or a very shal-
Uses
low groove on adaxial (upper) side, an obtuse keel
The wood of this species is considered to be of good on abaxial side. Stomata small, in intermittent white
quality for construction of houses. The pendulous or lines forming narrow bands on either side of abaxial
drooping habit of the foliage would make an attrac- midrib. Pollen cones axillary, solitary or with 2–3 on
tive feature for horticulture and provenances from slender, 3–4 mm long peduncles, with a few trian-
the highest altitudes may even prove more or less gular perular scales at their base, cylindrical, 8–15
hardy. It is not known to be in cultivation. mm long, 3 mm wide at anthesis; microsporophylls
with an elongated, rostrate apex, bearing two ovoid-­
globose pollen sacs at base. Seed cones axillary,
solitary on short peduncles; receptacles subtended
by two minute, deciduous bracts (‘foliola’), consist-
ing of a short axis with two unequal bracts which
become fused and coriaceous, 2–3 mm long. Seeds
solitary with fertile bract, including the epimatium
ca. 8 × 5 mm, obliquely ovoid with a minute crest.
 Podocarpus pilgeri Foxw., Philipp. J. Sci. 2: 259.
Taxonomic notes
1907. Podocarpus celebicus Warb., Monsunia 1:
In the herbarium, this species could be confused 192. 1900, non Hemsl. (1896). Types: Indonesia:
with Podocarpus rostratus. Gaussen (1976) empha- Sulawesi, Wawo-Kraeng, O. Warburg 16433, 16890
sised leaf anatomical characters as the distinction of (syntypes B, destroyed, duplicates may exist in
the two species: Podocarpus perrieri has only 3 resin other herbaria).
canals, while P. rostratus, with slightly larger and
wider or flatter leaves, has many. To observe this Podocarpus pilgeri Foxw. var. thailandensis Gaussen,
character, microtome-cut cross-sections of fresh Trav. Lab. Forest. Toulouse T. 2, 1 (2, 21): 185. 1976
leaves have to be prepared for microscopic observa- (nom. inval., Art. 37.1).
880 tion. Podocarpus tixieri Gaussen, Trav. Lab. Forest. Tou-
louse T. 2, 1 (2, 21): 155. 1976 (nom. inval., Art. 37.1);
Podocarpus tixieri Gaussen ex Silba, J. Int. Conifer
Distribution
Preserv. Soc. 15 (2): 34. 2008.
Madagascar: Fianarantsoa Province (Andringitra
Massif), Antanarivo Province (Toamasina, Forêt
Etymology
d’Andasibé).
TDWG codes: 29 MDG This species was named in honour of the German
botanist Robert Pilger (1876–1953).
Ecology
Vernacular names
Podocarpus perrieri occurs in the eastern subhu-
mid forest, at elevations between 1200 m and 2000 In western New Guinea (Papua) this species is
(–2500?) m a.s.l. In one of the two known locations known by the name bempop; several names are re-
the terrain is described as “pentes rocailleuses”, i.e. corded in eastern New Guinea (Papua New Guinea)
rocky slopes, at an altitude of 2000 m, and the rocks and listed in Flora Malesiana ser. 1, 10 (3): 410–411
are silicious. At the type locality, at 1200 m a.s.l., this (1988). In the Philippines it is called lubang-lubang
species was said to attain 30 m in height; this would (Manobo) and in Thailand phayamai-baisan.
have been in tall forest. No mention of tree size was
made with the collection made at 2000 m a.s.l.
Description
Decumbent or erect shrubs or medium size trees
Conservation
to 25 m tall; trunk of trees monopodial, erect, to 60
This species is only certainly known from the type cm d.b.h., sometimes fluted at base. Bark of trees fi-
location and one other location and from GIS data it brous, flaking in strips, brown weathering grey; in-
appears to be a deforested area. The type collection ner bark pink. Branches ascending or spreading, in
was made in 1925, the other collection dates from trees forming a narrow or spreading crown. Foliage
1951. Surveys in this area are urgently required to de- branchlets terete, slender, terminating in small buds
termine if this species is still extant. Deforestation is 3–4 mm long with erect or slightly spreading, tri-
serious in this region and this species is not known angular to lanceolate and narrowly acute scales up
from any protected area. to 4 mm long. Leaves on seedlings and young trees
IUCN: CR [B1, B2, a-b (iii)] and/or shaded branches of mature trees usually
larger than on shrubs and sun-exposed branches of
mature trees, to 7 cm long and 12 mm wide, taper-
Uses
ing at both ends. Leaves on shrubs and mature trees
No commercial uses are recorded of this species, but when sun-exposed (1.5–)2.5–4.5(–5) cm long, (4–)5–
it is undoubtedly used locally for firewood and, if 7(–10) mm wide, linear-elliptic to oblong, greatest
trees are large enough, for construction timber, such width usually above middle, gradually tapering to a
as flooring of houses. short petiolate, slightly twisted base; apex variable,
often obtuse to abruptly rounded, sometimes acute
or cuspidate especially in exposed leaves of shrubs; the crests and summit areas of Mt. Hunstein (Papua
margins slightly revolute. Midrib on adaxial (upper) New Guinea) and Mt. Nettoti (Papua, Arfak Mts.)
side 0.2–0.3 mm wide, acutely raised, on abaxial where Podocarpus pilgeri is a decumbent shrub. At
side wider, to 1 mm, obtuse or nearly flat. Leaf col- its highest point (3300 m) it grows on the edge of
our variable; upperside dark glossy green, dull green low, open ‘heath forest’ and subalpine grassland on
or glaucous; underside light green or grey-green. peaty soil.
Stomata very small, numerous, in two broad bands
on abaxial side. Pollen cones axillary, solitary, cylin-
Conservation
drical, 15–30 mm long, ca. 3 mm wide, with a few
rounded scales (from buds) on a 2–4 mm long pe- IUCN: LC
duncle; microsporophylls triangular-apiculate, with 881
two globose pollen sacs at base. Seed cones axillary,
Uses
solitary; receptacles with two 1.5–2 mm long basal
bracts (‘foliola’), 10–12 mm long and much swollen Where it grows as a medium-size forest tree it will
when ripe, greenish yellow, becoming bright red be logged and used in similar ways as other ‘podo-
or purple. Seeds including the epimatium ovoid- carp’ trees, i.e. for construction (house building) and
globose, 10–12 × 8–9 mm, smooth without a crest other carpentry. Its scattered occurrence and often
at maturity, glaucous green to purplish green. Seed shrubby habit make it less important in the tim-
proper ovoid-globose, 8–10 mm long; micropylar ber trade, so most uses will be rather local. It is not
end slightly elongated and curved. known to be in cultivation outside a few collections
in botanic gardens.
Distribution
S China: Hainan Island, Guangxi; Indochina: Kam- Podocarpus polyspermus de Laub., Fl. Nouv.
puchea [Cambodia], Laos, N Vietnam; Malesia: Calédonie Dépend. 4: 60. 1972. Type: New
Admiralty Islands, Borneo (Sarawak), Maluku Caledonia: Grande Terre, Province Sud, Mé Maoya
[Moluccas] (Obi), Philippines, Sulawesi; Papuasia: [Mt.], ridge north of the Col des Roussettes, D. J. de
New Guinea. Laubenfels P 424 (holotype P).
TDWG codes: 36 CHH CHS-GX 41 CBD LAO VIE 42
BOR-SR MOL PHI SUL 43 NWG-IJ NWG-PN
Etymology
The species epithet refers to the multiple seeds that
Ecology
are found on many receptacles, instead of the single
Podocarpus pilgeri occurs in montane forests (often seed in most species.
mossy forest) and in low scrub on exposed moun-
tain ridges and summits. Its altitudinal range is
Vernacular names
(700–)1200 to 3000(–3300) m a.s.l. This species is
in some localities a decumbent shrub not taller than No common names are known for this species.
1 m, elsewhere it attains tree sizes. Apparently this
is not correlated with altitude, since a tree 10 m tall
Description
was sampled at 3300 m on Mt. Wilhelm in Papua
New Guinea and low shrubs have been found at Small trees 5–15 m tall, monopodial, to 25 cm d.b.h.
around 1000 m altitude. It becomes dwarfed on ex- Bark fissured and scaly, dark or light brown; in-
posed mountain ridges and summits amongst rocks. ner bark slightly fibrous. Branches spreading and
Trees are often limited to only reaching subcanopy assurgent, forming an irregular and open crown.
height, in which case the forest is usually open, but Foliage branches terete, stout, ridged between leaf
it survives as well under canopy for a time. In New bases, with leaves mostly at distal parts, terminat-
Guinea it is often associated with Southern beech ing in obtuse conical buds surrounded by elon-
(Nothofagus sp.), while another common conifer in gated and spreading or reflexed outer scales 10–15
this ‘beech forest’ is Phyllocladus hypophyllus. Low mm long and 2 mm wide at base. Leaves on juve-
mossy Xanthomyrtus-Podocarpus forest dominates nile plants not known; on mature trees crowded,
 more or less erect or spreading at a small angle to
Uses
shoot, linear-­lanceolate, 5–12 cm long, 6–8 mm
wide, straight or slightly falcate, gradually tapering No uses have been recorded of this species.
to a petiolate base, coriaceous; apex acute or obtuse.
Midrib prominent on both sides, on adaxial (upper)
side obtuse, on abaxial side flattened but with sharp Podocarpus polystachyus R. Br. ex Endl., Syn.
sides. Leaf colour lustrous grey-green or dark green Conif.: 215. 1847. Margbensonia polystachya
above, dull green below, flushing glaucous green. (R. Br. ex Endl.) A. V. Bobrov & Melikyan, Bjull.
Stomata small, in numerous intermittent lines in Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1):
two broad bands on either side of abaxial midrib. 60. 1998. Type: Singapore: [“Legi in Singapore
882 Pollen cones axillary, solitary, sessile; from scaly, ad libra”], N. Wallich 6052B (lectotype K-W,
subglobose buds sometimes persisting after leaves designated here). Fig. 286
have fallen; bud scales triangular, keeled; cones
elongating to cylindrical, 20–25 mm long, 3.5–5 mm Podocarpus thevetiifolius Zipp. ex Blume, Rumphia
wide; microsporophylls spirally arranged, imbri- 3: 213. 1847 [“thevetiaefolius”]; Podocarpus poly­
cate, with an elongated apex and two basal pollen stachyus R. Br. ex Endl. var. thevetiifolius (Zipp. ex
sacs. Seed cones axillary, solitary on 5–10 mm long, Blume) Silba, Phytologia 68: 70. 1990; Margbenso­
stout peduncles; receptacles formed of an axis with nia thevetiifolia (Blume) A. V. Bobrov & Melikyan,
3–5 bracts subtended by two spreading or recurved Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1):
bracts (‘foliola’) 3–4 mm long, fusing and swelling to 60. 1998.
10–13 × 8–9 mm, becoming succulent and red. Seeds
often 2 per receptacle, sometimes 1 or 3, including
Etymology
the epimatium 8 × 6 mm, obovoid-globose without
a crest. Seed proper not observed. The species epithet means ‘with many spikes’ and
refers to the clustered pollen cones.
Distribution
Vernacular names
New Caledonia, on several mountains in the central
part of Grande Terre. podo laut (Malay); jati bukit (Peninsular Malaysia);
TDWG codes: 60 NWC landin (Sarawak); kandabang (Sabah) and several
more locally used names as listed in Flora Malesiana
ser. 1, 10 (3): 418 (1988).
Ecology
Podocarpus polyspermus has been found on moun-
Description
tains at altitudes between 650 m and 950 m a.s.l. in
low forest or transitions between ‘maquis minier’ Trees to 20(–40) m tall, d.b.h. to 45 cm (common-
and forest, on serpentine or ultramafic rocks de- ly a monopodial or multi-stemmed small tree ca.
rived from this. On mountain ridges the forest is 6 m tall). Bark shallowly fissured, peeling in long,
often open and low and this species forms a canopy fibrous strips, brown weathering grey; inner bark
tree with a maximum height of 5–7 m and a charac- soft fibrous, pink. Branches spreading, forming an
teristic flat candelabra crown; on less exposed sites irregular to rounded and dense crown in most trees.
it remains below the canopy formed often by an- Foliage branchlets terete, slightly grooved or ridged,
giosperms, with emergent Araucaria spp. terminating in 1.5–3 mm long buds with narrowly
triangular to lanceolate, erect to spreading outer
scales. Leaves on juvenile plants linear to linear-­
Conservation
lanceolate, 7–11 cm long, 8–13 mm wide, more or less
In several parts of its presently imperfectly known abruptly tapering to a petiolate base and more grad-
range this species is under threat by nickel mines ually to an acute or apiculate apex. Leaves on mature
and their expansion. Only one population is in a trees linear to linear-lanceolate or nearly oval when
protected area. very short, (2.5–)5–11 cm long, 6–11 mm wide, short
IUCN: EN [B1ab(iii)+2ab(iii)] petiolate, abruptly narrowed at base, more ­gradually
tapering to an acute or obtuse apex, straight or only occur in competition with other tall forest trees,
slightly curved, coriaceous; margins flat or nearly so; where P. polystachyus is striving to reach the canopy.
midrib acutely raised and narrow adaxially (above), It appears therefore to be a highly adaptable species
only slightly raised and wider abaxially (below); leaf and the nearest to a mangrove-inhabiting conifer in
colour deep green above, pale green below. Stomata existence.
on abaxial side, very small and in numerous irregu-
lar lines in two broad bands divided by midrib. Pol-
Conservation
len cones clustered axillary with 2–5, sessile or very
short pedunculate, with a few small scales at base, IUCN: LC
cylindrical to 15–30 mm long and 3 mm wide; mi-
crosporophylls imbricate, slightly spreading at an- 883
Uses
thesis, apiculate, with two oblong pollen sacs. Seed
cones axillary, solitary on a 2–6 mm long peduncle; Where this species grows into a tall tree, e.g. in
receptacles with 2 very small basal bracts to 1.5 mm Maluku [Moluccas], it is a fairly important source
long (deciduous), 7–10 mm long, swelling to 8 mm of ‘podocarp’ timber. It has pale yellowish brown,
thick, pruinose, becoming orange-red to red when leight-weight wood used for construction, window
ripe. Seeds including the epimatium 7–9 × 5–7 mm, frames, boat building including masts, spars and
globose-ovoid, smooth, glaucous green. Seed proper oars, flooring, veneer, furniture making and cabi-
not observed. net work, carpentry and joinery, household utensils,
matches, and toothpicks. Podocarpus polystachyus is
one of few species in the genus commonly planted
Distribution
as an ornamental tree in tropical countries, mostly
Indochina: southernmost peninsular Thailand; within its area of natural distribution but at least as
Malesia: Borneo, Peninsular Malaysia, Maluku far away as the Congo Republic.
[Moluccas] (Waigeo Island), Philippines, Singapore,
Sulawesi, Sumatera (Bangka & Belitung Islands);
­Papuasia: New Guinea (Papua: Vogelkop Peninsula). Podocarpus pseudobracteatus de Laub., Blumea 26
TDWG codes: 41 THA 42 BOR-BR BOR-KA BOR- (1): 142. 1980. Type: Papua New Guinea: Southern
SB BOR-SR MLY-PM MLY-SI MOL PHI SUL SUM 43 Highlands, Tari Subdistrict, Mt. Ambua, C. Kalk­
NWG-IJ man 5189 (holotype L).

Podocarpus archboldii N. E. Gray var. crassiramosus

Ecology
N. E. Gray, J. Arnold Arbor. 39: 453. 1958.
This species occurs primarily on sandy beaches and
bluffs, often on the land side of mangrove thickets at
Etymology
or just above the high tide mark. It can even occur
within the mangroves on slightly raised, sandy ridg- The species epithet means ‘seemingly but not actu-
es (probably old beaches). Here it is a stunted tree ally like bracteatus’ and refers to the similar species
not exceeding 6–7 m in height. On coastal limestone P. bracteatus Blume from Jawa, the Lesser Sunda Is-
and granitic rocks the trees are gnarled and shrubby, lands and Sumatera.
not exceeding 2–3 m. A second important habitat
is coastal lowland kerangas, a low forest on almost
Vernacular names
pure sand, and ‘pandangs’, i.e. degraded heath for-
est; in these vegetations Myrtaceae have an impor- kaip, kebu, puling (Papua New Guinea)
tant presence besides the conifers. In the interior of
Peninsular Malaysia, the Philippines, and elsewhere
Description
it occurs on limestone hills and plateaus at altitudes
between 150 m and 550 m (in Palawan at ca. 1000 Trees to 15(–20) m tall, but usually smaller; trunk
m) a.s.l. Here trees may be taller; in Obi (Moluccas) short, to 40 cm d.b.h. Bark fibrous, becoming flaky
trees are reported to reach 40 m tall with slender, on largest stems, peeling in longitudinal strips, cinna-
buttressed boles clear of branches to 25 m. This can mon brown or dark brown. Crown usually ­irregular
 or stunted. Foliage branches spreading, mostly with
Uses
widely spaced leaves. Terminal buds on leading
shoots 6–14 mm long, 3–4 mm wide, narrowly coni- Being usually a stunted tree, this species has no com-
cal, with free, long acuminate outer scales. Leaves on mercial value for timber; other uses are not known.
young plants 15–22 cm long, 12–17 mm wide, linear-
lanceolate, straight or slightly curved, acute. Leaves
on mature plants shorter and narrower, (4–)6–15 cm Podocarpus purdieanus Hook., Icon. Pl. 7: t. 624.
long, (5–)7–11 mm wide, lanceolate to linear-lanceo- 1844. Type: Jamaica: [“woods on the estate of
late, the shortest leaves elliptic, more or less abruptly Dunrobin Castle, … 8 miles north of White Hall”],
tapering to a petiolate base, gradually tapering to an W. Purdie s.n. (holotype K).
884 acute apex; margins flat; midrib acutely raised and
less than 0.5 mm wide on adaxial side, obtuse and
Etymology
ca. 1 mm wide on abaxial (lower) side. Stomata very
small and inconspicuous in dense lines on abaxial This species was named by William Hooker after
side. Pollen cones axillary, solitary or occasionally in W. Purdie, who collected the first known specimens
pairs, short pedunculate, subtended by short obtuse in 1843–44.
bracts (bud scales), elongating at anthesis to 30–45
mm, 3–4 mm wide; microsporophylls with acute
Vernacular names
apex ca. 1 mm long, bearing two globose pollen sacs.
Seed cones axillary on a 2–4 mm long peduncle; re- No common name has been recorded for this spe-
ceptacles subtended by a 3 mm long, thick bract (‘fo- cies.
liolum’), 8–11 mm long, formed of an axis with two
unequal bracts, swelling to become succulent and
Description
first orange, then red. Seeds including the epima-
tium ovoid-globose, 10–11 × 8–9 mm when mature, Trees to 40 m tall or more, monopodial, erect; trunk
distally obtuse. Seed proper not observed. d.b.h. to 1 m. Bark thin, smooth becoming scaly with
longitudinal strips, brown weathering grey. Branch-
es spreading, forming a rounded crown in large ma-
Distribution
ture trees. Foliage branches slender or stout, terete,
New Guinea. longitudinally grooved between leaf bases, termi-
TDWG codes: 43 NWG-IJ NWG-PN nating in more or less ovoid buds 3–4 mm wide
with imbricate, thick, strongly acuminate scales
with scarious thin margins and free apices. Leaves
Ecology
on saplings and vigorous shoots up to 14 cm long
Podocarpus pseudobracteatus is a species of montane and 17 mm wide. Leaves on mature trees 4–7(–9) cm
to high montane mossy Castanopsis-Nothofagus for- long, (6–)8–12(–14) mm wide, linear-lanceolate to
est, where it is commonly a canopy tree. At higher sub-spathulate (widest above the middle), straight
altitudes and on exposed ridges and summits, this or nearly straight, coriaceous, lustrous green above,
forest becomes low and stunted and trees attain only pale green below; margins flat or slightly revolute,
5–10 m, but in more protected sites where the for- narrowing gradually to a short petiolate base and
est grows taller it may attain larger size. Its altitudi- gradually or more abruptly to an acute, apiculate or
nal range is from 1700 m at the lowest limit to more apiculate-pungent apex. Midrib on adaxial (upper)
commonly 2200–2850 m a.s.l. At high altitudes this side raised only near base in most leaves, often in a
species can occur in swamp forest with Dacrycarpus shallow groove, continuous to apex but flat or with
and Dacrydium, or it may enter the alpine shrub- a central groove (in sicco) on abaxial side. Stomata
bery dominated by Ericaceae, Myrtaceae, and some- very small, in numerous intermittent lines on abax-
times Podocarpaceae. ial surface. Pollen cones axillary, solitary, sessile, cy-
lindrical, 15 mm or more long (fully elongated cones
not observed); microsporophylls more or less ovate,
Conservation
obtuse, bearing two basal pollen sacs. Seed cones
IUCN: LC axillary, on 2–4 mm long peduncles; ­receptacles
with two deciduous basal bracts (‘foliola’) and with c­ onstruction, floors, furniture making, and numer-
spreading bract tips apically, swelling to a succulent, ous other uses by the European settlers. Large trees
red, 7–8 mm long body (pseudo-fruit). Seeds ob- in accessible places became scarce and logging di-
liquely ovoid, including the epimatium 8 × 5 mm, minished to occasional instances. More recently,
with a small conical, obtuse crest. Seed proper not saplings of this and other tree species are being cut
observed. on a large scale as sticks (props) for the cultivation of
yams (Dio­scorea sp.), a tuberous twining plant and a
staple food. This species is not known in cultivation.
Distribution
Caribbean: Jamaica (Mt. Diablo, Sander’s Hill).
TDWG codes: 81 JAM Podocarpus ramosii R. R. Mill, Edinburgh J. Bot. 885
63 (1): 81. 2006. Podocarpus rotundus de Laub., Ka-
likasan 7 (2): 136. 1978 (nom. illeg., Art. 53.1). Type:
Ecology
Philippines: Luzon, Laguna Prov., Mt. Banáhao,
Podocarpus purdieanus occurs in wet tropical ev- M. Ramos BS 19581 (holotype US).
ergreen forest on limestone. The elevation is 800
m a.s.l. at the Mt. Diablo subpopulation; the range
Etymology
given in the literature is 720–1050 m a.s.l.
The species epithet commemorates Maximo Ramos
(d. 1932), who made extensive plant collections in
Conservation
the Philippines.
A region measuring 60 km × 30 km is the area
where it might be found, giving an extent of occur-
Vernacular names
rence (EOO) of 1800 km2. Mt. Diablo has the best
known sub-population, but it may be little more No common names are recorded for this species.
than 10 mature trees. Another sub-population, on
Sander’s Hill, has 15 trees found in dense forest,
Description
with many seedlings and saplings seen. There are
probably other small subpopulations in the area. At Small trees to 15 m tall; trunk monopodial, erect, to
present, only two locations are known; in the past it 30 cm d.b.h. Bark smooth, becoming scaly, brown
has been collected near Moneague, Troy in Trelawny weathering grey; inner bark pink. Branches spread-
Parish, and elsewhere. In the 19th century it was said ing, forming a spreading crown. Foliage branchlets
to grow in “woods on mountain ridges” and to be terete, slender, finely grooved, terminating in small
“one of the noblest trees in the island.” Historically, buds 2–3 mm long with erect or slightly spreading,
it was probably used for rebuilding ships; some log- triangular to lanceolate and narrowly acute scales
ging continues. Present day threats are deforestation up to 3 mm long. Leaves on seedlings and young
and cutting saplings for yam sticks. One million trees and/or shaded branches of mature trees usu-
yam sticks are required per year, not just made from ally larger than on sun-exposed branches of mature
Podocarpus, but saplings and small trees are taken trees, to 7(–10?) cm long and 15 mm wide, linear-
from the forest where P. purdieanus grows. There is lanceolate; apex obtuse to acute. Leaves on mature
no legislation that regulates the logging or cutting trees when sun-exposed (1.5–)2.5–5 cm long, 7–13
of this species and it has not been found to occur mm wide, elliptic to obovate, greatest width usu-
within any protected area. ally above middle, gradually or abruptly tapering
IUCN: EN [B1, a-b (ii, iii, v)] to a short petiolate base; apex obtuse to abruptly
rounded; margins slightly revolute. Midrib on adax-
ial (upper) side 0.2–0.3 mm wide, acutely raised, on
Uses
abaxial side wider, to 1 mm, obtuse or nearly flat.
Trees of large size of this species have been logged Leaf colour on upperside dark glossy green, on un-
extensively in the past. The wood of this species derside light green. Stomata very small, in intermit-
is more highly prized for timber than mahoga- tent, wavy lines in two broad bands on abaxial side.
ny. The wood was used for ship building, house Pollen cones axillary, solitary, cylindrical, 20–30
 mm long, ca. 3 mm wide, with a few lanceolate to
Conservation
linear scale leaves on a 5–10 mm long peduncle; mi-
crosporophylls less than 1 mm long, with two large Podocarpus ramosii is only known from two very
globose pollen sacs at base. Seed cones axillary, soli- disjunct populations. We are unsure of the threats
tary, on a 5–10 mm long peduncle; receptacles with to the habitat in either of these and neither of them
two 2 mm long basal bracts (‘foliola’), 7–10 mm long are within protected areas. They will probably not be
and swollen when ripe, greenish yellow, becoming subject to logging or deliberate deforestation due to
red. Seeds including the epimatium ovoid-globose, respectively small sizes of trees and unsuitability of
8–10 × 6–7 mm, smooth without a crest at maturity, the localities for agriculture. The conservation sta-
glaucous green. Seed proper not observed. tus of this species is therefore still unknown. More
886 information is also needed to confirm (or reject) its
status as a distinct species.
Taxonomic notes
IUCN: DD
There is some doubt about the taxonomic distinc-
tiveness of this species. The type collection is from
Uses
Mt. Banahao in Luzon, Philippines, as are most
other gatherings; only one sampling (W. Meijer 903) No uses have been recorded of this species.
was taken from Mt. Beratus in Borneo. The most
similar species is P. pilgeri, which mainly differs in
having narrower and more pointed leaves at least Podocarpus ridleyi (Wasscher) N. E. Gray, J. Arnold
in sun-exposed branches. De Laubenfels (op. cit.) Arbor. 39: 435. 1958. Podocarpus neriifolius D. Don
stated, when he described this species with the il- var. ridleyi Wasscher, Blumea 4 (3): 453. 1941;
legitimate name P. rotundus (a later homonym), that Margbensonia ridleyi (Wasscher) A. V. Bobrov &
P. pilgeri had not been found in Borneo or in Luzon. Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
However, it has been collected in Sarawak as well as Biol. 103 (1): 60. 1998. Type: Malaysia: Peninsular
in NW Luzon with specimens that, based on their Malaysia, Johore, Gunung Ledang [Mt. Ophir],
morphological characters, could not be included H. N. Ridley 10016 (holotype S).
with P. ramosii (P. rotundus) without making the lat-
ter synonymous to P. pilgeri. Perhaps P. ramosii is a
Etymology
subspecies or variety of P. pilgeri with broad, apically
rounded leaves. Podocarpus costalis, another name This species commemorates the plant collector
given to some specimens of P. ramosii, is indeed a H. N. Ridley, who collected the type specimen.
very different species and occurs in a very different
habitat as well.
Vernacular names
No vernacular names are recorded for this species,
Distribution
which was originally described as a mere variety of
Indonesia: Kalimantan Timur (Mt. Beratus); Philip- the widespread P. neriifolius.
pines: Luzon (Mt. Banahao).
TDWG codes: 42 BOR-KA PHI
Description
Trees to 25 m tall, in stunted forests less than 10 m
Ecology
tall, d.b.h. to 40 cm. Bark soft, flaky or scaly, pale
Podocarpus ramosii occurs in dwarf mossy forest, brown; inner bark dark red-brown. Branches
ant an altitude of ca. 1000 m a.s.l. on Mt. Beratus, up spreading, forming an irregular crown in most trees.
to ca. 2100 m on Mt. Banahao in Luzon. Mt. Beratus Foliage branchlets terete, slightly grooved or ridged,
is only 1223 m and Mt. Banahao in Luzon peaks at terminating in 4–8 mm long buds with narrowly
2170 m, so in both localities this species occurs near triangular to lanceolate, erect to spreading scales.
the summit. The geographical disjunction may be Leaves on juvenile plants linear-lanceolate, 11–20 cm
real, but could well be an artifact of poor collecting, long, 10–16 mm wide, gradually tapering to a peti-
or poor recognition as a distinct species. olate base and to a narrowly acute apex. Leaves on
mature trees lanceolate to linear-lanceolate, (4–)6– Podocarpus roraimae Pilg., Notizbl. Bot. Gart.
12(–14) cm long, 8–15 mm wide, distinctly petiolate, Berlin-Dahlem 5: 299. 1913. Type: Guyana: Cuyuni
gradually narrowed at base, very gradually tapering Mazaruni, Pakaraima Mountains, Mt. Roraima,
to a narrowly acute apex, straight or slightly curved, E. H. G. Ule s.n. (holotype B).
coriaceous; margins somewhat revolute; midrib in-
distinctly raised and narrow adaxially (above), more Podocarpus buchholzii de Laub., Fl. Venezuela 11 (2):
prominently raised and wider abaxially (below); 31. 1982.
leaf colour yellowish green above, pale green below. Podocarpus buchholzii de Laub. var. neblinensis
Stomata on abaxial side, very small and in numer- Silba, Phytologia 68: 66. 1990.
ous irregular lines in two broad bands divided by
midrib. Pollen cones clustered axillary with (1–)2–4 887
Etymology
or several on small lateral leafless shoots, sessile or
very short pedunculate, with papery scales at base, The species epithet refers to Mt. Roraima, on the
cylindrical to 15–20 mm long and 3 mm wide; mi- border between Guiana and Venezuela.
crosporophylls imbricate, slightly spreading at an-
thesis, apiculate, with two oblong pollen sacs. Seed
Vernacular names
cones axillary, solitary on a 3–12 mm long peduncle;
receptacles with 2 small basal bracts (‘foliola’) to 2.5 Ai-yek (Venezuela)
mm long (sometimes lacking in early stages, so not
deciduous), 8–10 mm long, swelling to 8 mm thick,
Description
becoming pink when ripe. Seeds including the epi-
matium 7–8 × 4–5 mm, obliquely ovoid, without a Shrubs or small trees 1–3 m or 7–10 m tall; trunk
crest, glaucous. Seed proper not observed. diam. to 30 cm. Branches spreading, in older plants
forming a dense, often straggling crown. Foliage
branchlets slender or more robust, terete, finely
Distribution
grooved between leaf bases, terminating in sub-
Malaysia: Peninsular Malaysia. globose buds 3–5 mm wide, with free, erect, acute
TDWG codes: 42 MLY-PM scales, the outer scales often transformed into or ac-
companied by reduced leaflets (much) longer than
normal bud scales. Leaves on vigorous shoots of
Ecology
juvenile plants remotely placed, spreading, on ex-
Podocarpus ridleyi occurs localized on several posed, slower growing branches crowded, spreading
mountain summits and ridges in somewhat stunted at narrow angles to shoot to nearly erect, oblan-
rain forest; it may attain greater size when it occurs ceolate (widest beyond the middle) to elliptic, on
in forest on slopes below these summits. Its altitudi- juvenile plants somewhat linear and to 6 cm long,
nal range is from 480 m to 1500 m (possibly to 2100 on mature plants 2–3(–3.8) cm long, 4–7 mm wide,
m) a.s.l. It grows in impoverished soils derived from gradually narrowing to a short petiolate base, more
sandstone or granite (Mt. Ophir) and can be the abruptly ending in an acute or obtuse apex. Margins
dominant tree in these localities. revolute; leaf texture coriaceous; leaf colour deep
green above, pale or glaucous green below. Midrib
on adaxial (upper) side obtusely raised or nearly
Conservation
flat with a central groove, on abaxial side promi-
IUCN: LC nently raised. Stomata on abaxial side very small,
in numerous intermittent lines on either side of the
midrib. Pollen cones axillary, solitary, sessile with
Uses
perular scales at base, cylindrical, 15–20 mm long,
No uses have been recorded of this species. 2.5–3 mm wide; microsporophylls triangular, with
two globose pollen sacs. Seed cones axillary, solitary
on a 5–10 mm long, slender peduncle; receptacles
small, 6–8 mm long, formed of a short axis with
two unequal bracts, slightly swelling and ripening
 red. Seeds ­single, including the epimatium ca. 8 mm
Conservation
long, ca. 5 mm wide, ovoid with a distal crest. Seed
proper not observed. This species, originally described from Mt. Roraima,
has since been found in several isolated tepuis in
Bolivar and Amazonas, Venezuela. Due to the re-
Taxonomic notes
moteness of this region and its mostly undisturbed
Podocarpus buchholzii was described as a distinct montane rainforests, it is believed that this species
species by David de Laubenfels (op. cit.) differing may occur on yet other mountains and is not threat-
from P. roraimae in its leaf midrib, described as a ened. One known location is within Roraima Na-
groove on the adaxial side, instead of having a raised tional Park.
888 midrib as described in P. roraimae. The specimens IUCN: LC
actually show variation in this character, with many
leaves having a slightly raised midrib with a groove
Uses
in the middle, others have a raised midrib from
the leaf base which turns into a grooved midrib to- No uses have been recorded of this species.
wards the apex and still others have a groove on a
flat midrib along the entire leaf length. In all other
morphological characters, the two taxa are entirely Podocarpus rostratus L. Laurent, Ann. Fac. Sci.
similar and they occur in similar habitats, from bog Marseille 23: 60. 1915. Type not designated.
margins to low woodland in ravines. It is therefore
considered that P. buchholzii, and the variety under
Etymology
that name described by John Silba (op. cit.), are taxo-
nomic synonyms of P. roraimae. The species epithet (Latin rostrum = beak) means
‘provided with a beak’ and refers to the apical shape
of the microsporophyll.
Distribution
Guyana (Mt. Roraima); Venezuela (several isolated
Vernacular names
mountains in Amazonas and Bolivar).
TDWG codes: 82 guy VEN No common names have been recorded for this spe-
cies.
Ecology
Description
Podocarpus roraimae is found on isolated table
mountains (tepuis), mostly on sandstone, at alti- Small, usually stunted trees 8–10 m tall. Bark ex­
tudes between 1800 m and 2700 m a.s.l. This spe- foliating in small strips and flakes, light brown
cies occurs in small stands of shrubs and trees in a weathering grey. Foliage branchlets slender, terete,
mozaic vegetation of ombrotrophic bogs dominated finely grooved on vigorous shoots of young plants,
by grasses and herbs and interspersed low scrub and ridged between decurrent leaf bases on older, slow
woodland. It also occurs in ravines near streams and growing trees, terminating in small ovoid-globose
on wet plateaux and summits with boggy vegetation buds with imbricate, apiculate scales. Leaves on
characterized by various ferns, pitcher plants and young plants linear, 2.5–6 cm long, 2–3 mm wide,
orchids. Shrubs of this species occur in more open straight or curved downward. Leaves on mature
terrain, while in forest it can attain small tree size. It trees narrowly lanceolate to linear, (1–)1.5–3(–3.6)
can be associated with P. steyermarkii and P. tepuien­ cm long, straight or sometimes slightly falcate to-
sis, which are distinct in having larger, respectively wards apex, 1–2 mm wide, only slightly narrowing
smaller, leaves with a midvein groove on the adaxial to a sessile, decurrent base; margins entire, mostly
side. There is high rainfall and frequent mist, and parallel but in the distal part tapering to an acute
temperatures can fluctuate strongly from day to apex. Midrib absent or inconspicuous on adaxial
night and depending on weather conditions. (upper) side by a shallow central groove, continuous
but flat or obtusely raised on abaxial side. Stomata
Uses
on abaxial side in several intermittent lines on ­either
side of midrib. Pollen cones axillary, solitary or in No uses have been recorded of this species. It is not
pairs, rarely 3 on 1–6 mm long stalks, subtended known to be in cultivation, but should be attempted
by a few keeled bud scales, cylindrical, 10–15(–20) in the context of ex situ conservation to back-up the
mm long, 2.5–3 mm wide; microsporophylls more remaining fragmented populations in the wild.
or less triangular, with an elongated, 1 mm long ros-
trate apex and two relatively large basal pollen sacs.
Seed cones axillary, solitary, pedunculate, formed of Podocarpus rubens de Laub., Blumea 30 (2): 266.
an axis with two unequal bracts, subtended by two 1985. Type: Papua New Guinea: Western Highlands,
spreading or recurved 1 mm long bracts (‘foliola’), Wahgi-Sepik Divide, J. S. Womersley & A. N. Millar 889
fusing and slightly enlarging to a cuneate, 3–4 × 2–4 NGF 6980 (holotype L).
mm, coriaceous receptacle with divergent bract tips.
Seeds solitary, including the epimatium ca. 14 × 8 Podocarpus neriifolius D. Don var. timorensis Was­
mm, obliquely ovoid, with a small crest slightly be- scher, Blumea 4: 451. 1941.
low the apparent apex. Seed proper not observed. Podocarpus indonesiensis de Laub. & Silba, Phytolo-
gia 64: 292. 1988.
Podocarpus rubens de Laub. var. pabinamaensis
Distribution
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 37. 2000.
Madagascar: Antsiranana Prov., Fianarantsoa Prov., Podocarpus rubens de Laub. var. sumatrana Silba,
Mahajanga Prov., Toamasina Prov. J. Int. Conifer Preserv. Soc. 7 (1): 37. 2000.
TDWG codes: 29 MDG
Etymology
Ecology
The species epithet means ‘red’ and refers to the col-
Podocarpus rostratus is recorded from rocky slopes our of the flushing leaves.
on silicious rocks; the elevation is given as 1800–
2400 m a.s.l. (herbarium data), but the range may be
Vernacular names
greater according to GIS calculations and informa-
tion from literature. It occurs in ericaceous scrub- A number of local names have been recorded for
land near the summits of mountains, a vegetation New Guinea in Flora Malesiana, ser. 1, 10 (3): 402
type prone to fires. It is not well adapted to fires and (1988); in Indonesia this species is probably not
therefore occupies either open, rocky places with lit- distinguished by non-botanists and known under
tle vegetation, or wetter sites along streams. names used for more common, earlier described
species.
Conservation
Description
This species is known from three highly disjunct
populations. The greater number of herbarium col- Trees to 45 m tall, to 80 cm d.b.h., bole erect, straight.
lections is from the northern population and only Bark smooth, thin, exfoliating in long strips, pale
one from each of the eastern and southern popula- brown weathering grey; inner bark reddish brown,
tions (to over 1000 km away), made in 1924–25 at fibrous. Branches spreading, forming a rounded or
locations which have probably been deforested. All domed crown. Foliage branchlets terete, more or
but one collection are older than 50 years. The area less grooved, glabrous, terminating in 2–4 mm long
of occupancy (AOO) of the northern population is buds with erect or slightly spreading, triangular to
less than 500 km2 and only one sub-population is broadly lanceolate outer scales. Leaves on juvenile
inside a protected area, a collection was made there plants short petiolate, linear-lanceolate, 6–8 cm
recently (2005). Serious decline of small populations long, 10–14 mm wide, straight or slightly curved,
is therefore inferred for this species, which is sus- gradually narrowed at base, acuminate or apiculate.
ceptible to fires. Leaves of mature trees usually shorter and narrower,
IUCN: EN [B1, a-b (iii)] linear-lanceolate or elliptical, 3–6 cm long, 5–8(–10)
 mm wide, straight or slightly curved, short peti- growing conditions between that altitude and 1500
olate at a gradually narrowing base, tapering to an m. At higher altitudes it reappears on exposed ridg-
acute apex. Midrib acutely raised on both sides, usu- es between 2000 m and 3000 m a.s.l. as a small, of-
ally narrower on adaxial side; leaf colour dark green ten stunted tree in mossy forest, especially in New
above, pale green below; new leaves flushing bronze Guinea accompanied by Nothofagus spp. and other
or bright red. Stomata very small, in numerous ir- fagaceous trees. It is much less common in swamp
regular lines on abaxial (under) side on either side forest, where it may grow as an occasional tree with
of midrib. Pollen cones axillary, solitary or in clus- Dacrydium spp.
ters of 2–3, sessile or short pedunculate with several
basal bract scales, cylindrical, elongating to 20–35
Conservation
890 mm, 2.5–3.5 mm wide; microsporophylls triangu-
lar, spreading, with two globose pollen sacs. Seed IUCN: LC
cones axillary, solitary on a 4–9 mm long peduncle;
receptacles with 2 bracts (‘foliola’) 1–2 mm long at
Uses
base, obliquely bilobate, 6–8 mm long, swelling with
a truncate distal end, yellow-green turning red or Where it grows to a large tree, this species will be
purple and succulent when ripe. Seeds enclosed in valued and exploited for its timber. It has pale yel-
a smooth epimatium, green turning red or purple lowish brown, leightweight wood used for construc-
when ripe, ovoid, 8–9 × 5–6 mm, distal end obtuse. tion, window frames, boat building including masts,
Seed proper not observed. spars and oars, interior uses such as flooring, veneer,
furniture making and cabinet work, carpentry and
joinery, household utensils, matches, and tooth-
Taxonomic notes
picks. It is not known to be in cultivation.
The species P. indonesiensis de Laub. & Silba (op.
cit.), based on a few collections by P. J. Eyma in Sula­
wesi and an additional specimen from Amboina Podocarpus rumphii Blume, Rumphia 3: 214. 1847.
(Maluku), is perhaps only a high altitude form of Margbensonia rumphii (Blume) A. V. Bobrov &
P. rubens in Sulawesi (2500–3000 m a.s.l.) with short- Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
er and thereby more elliptical leaves. In New Guinea Biol. 103 (1): 60. 1998. Type: Indonesia: Papua
P. rubens has been collected from as high as 2750 m., [“Nov. Guinea”], leg. ign. s.n. (holotype L, barcode
a similar altitude as in Sulawesi. Shorter leaves tend L00050854). Fig. 287
to be elliptical, longer leaves linear-lanceolate, and
both occur on the same specimens (J. F. Veldkamp Podocarpus koordersii Pilg. ex Koord. & Valeton,
8318, K, L). The specimen from Amboina was said to Meded. Lands Plantentuin 68: 268. 1904; Margben­
be from around 1000 m a.s.l. All other characters are sonia koordersii (Pilg. ex Koord. & Valeton) A. V. Bo-
identical with P. rubens and P. indonesiensis is here brov & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
considered a taxonomic synonym. Otd. Biol. 103 (1): 60. 1998.
Podocarpus philippinensis Foxw., Philipp. J. Sci.
6: 149. 1911; Margbensonia philippinense (Foxw.)
Distribution
A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc.
Malesia: Borneo, Flores, Maluku [Moluccas] (Ser- Isp. Prir., Otd. Biol. 103 (1): 60. 1998.
am), Sulawesi, Sumatera, Timor; Papuasia: New Podocarpus rumphii Blume var. arbainii Silba, J. Int.
Britain, New Guinea. Conifer Preserv. Soc. 7 (1): 38. 2000.
TDWG codes: 42 BOR-KA BOR-SB BOR-SR LSI-LS Podocarpus rumphii Blume var. aruensis Silba, J. Int.
MOL SUL SUM 43 BIS NWG-IJ NWG-PN Conifer Preserv. Soc. 7 (1): 38. 2000.

Ecology Etymology
Podocarpus rubens is a constituent of lower montane This species was named in honour of Georg Eber-
to high montane primary rain forests. Its lowest al- hard Rumphius (1628–1702), author of an early Flora
titude is around 800 m a.s.l. and it reaches ­optimum of Amboina.
 Timor), Philippines (Luzon), Maluku [Moluccas]
Vernacular names
(Aru, Obi, Weda); Papuasia: New Guinea (including
Numerous local names are in use for this species; Misool and Numfoor Islands).
several are listed by region in Flora Malesiana ser. 1, TDWG codes: 36 CHH 42 BOR-SB JAW LSI-ET LSI-
10 (3): 416 (1988); in Vanuatu it is called nimsal (Sie LS MLY-PM MOL PHI SUL 43 NWG-IJ NWG-PN
language).
Ecology
Description
Podocarpus rumphii is a constituent of lowland to
Trees to 45 m tall, to 80 cm d.b.h., bole straight, in lower montane tropical rainforests, where it can be
forest clear of branches 10–20 m. Bark scaly, whitish locally common. It is likely to be confused with the 891
brown or brown; inner bark pinkish to red-brown. similarly widespread but more ubiquitous species
Crown spreading, rounded or domed and fairly P. neriifolius, which can have similarly large leaves
dense. Foliage branchlets terete, finely grooved, ter- (but usually narrower) and differs in characters not
minating in a globose to (short) conical bud (2.5–) always observable in specimens. The altitude ranges
4–5(–8) mm long, outer scales imbricate, meeting at from sea level to 1600 m, but most collections are
the papex, triangular, only at the tip sometimes bent from below 500 m a.s.l. In Jawa it is found on lime-
outward. Leaves on juvenile plants 19–26 cm long, stone, on other islands also on soils ranging from
18–25 mm wide, broad linear, straight; apex acute or clay to sand derived from acidic rock types. In tall
slightly acuminate. Leaves on adult trees on lower forest (40–50 m) it can reach into the canopy, suc-
branches or in shade 12–22 cm long, 10–20 mm cessfully competing with angiosperms. The large
wide, linear, straight or slightly curved, (long) peti- leaves of saplings to pole stage trees could be an ad-
olate and slightly twisted at base, gradually tapering aptation to growing in shade under other trees; they
towards base and towards acute apex, coriaceous become smaller, narrower and more leathery and
and stiff; leaves in exposed crown parts of large trees stiff in sun-exposed crowns of larger trees.
still shorter and narrower, 9–14 cm long, 10–15 mm
wide. Midrib on adaxial (upper) side conspicuous
Conservation
from base to apex, obtuse, ca. 1 mm wide, often col-
lapsing in dry leaves, midrib on abaxial side promi- Exploitation has been extensive and in the Philip-
nently raised. Stomata on abaxial side in numerous pines this has reportedly led to the situation that
lines on either side of midrib, small and inconspicu- this easily exploited tree survives over much of its
ous. Pollen cones axillary, in clusters of (2–)3–5, range only in protected spots. Due to its vast extent
sessile, subtended by imbricate, papery scales, long of occurrence (EOO) and the existence of numer-
cylindrical, 35–45 mm long, 4 mm wide at anthesis; ous localities where P. rumphii remains undisturbed,
microsporophylls initially imbricate, then parting, this species does not meet any of the criteria for a
small and apiculate, with two large, oblong pollen threatened category.
sacs. Seed cones axillary, solitary on a short pedun- IUCN: LC
cle; receptacles subtended by 2 minute bracts (‘fo-
liola’) 1–1.5 mm long, bilobed, with 3 lateral bracts,
Uses
one of which is smaller, ca. 10 mm long, swelling at
maturity to 20–23 × 9–11 mm, turning yellow then Podocarpus rumphii is a valuable timber tree where
red to purple. Seeds including the epimatium glau- it attains large sizes with a clear, straight bole. Its
cous or pruinose, turning blackish at maturity, sub- wood is used as roundwood for masts, spars, and
globose, 12–15 × 10–12 mm, with a curved distal crest poles, in house construction as beams, in high-
or beak. Seed proper subglobose, with a hard and grade construction for flooring, joinery and other
smooth seed coat, ca. 10 mm long. carpentry, for furniture and cabinet work, veneer,
to make boxes, and for match sticks. In traditional
use it was sought after for (dugout) canoes, used in
Distribution
coastal house construction, for household utensils,
China: Hainan Island; Malesia: Peninsular Malaysia, and wood carving. It is not known to be in culti-
Sabah, Jawa, Sulawesi, Lesser Sunda Islands (Flores, vation, either as a forestry plantation tree or as an
 ­ rnamental tree; the species is present in a few tropi-
o
Distribution
cal botanic gardens and arboreta.
Bolivia: Cochabamba, La Paz, Santa Cruz; Peru:
Cusco (Machu Pichu).
Podocarpus rusbyi J. T. Buchholz & N. E. Gray,
TDWG codes: 83 BOL PER
J. Arnold Arbor. 29: 134. 1948. Type: Bolivia: La Paz,
H. H. Rusby 2463 (holotype NY).
Ecology
Etymology
Podocarpus rusbyi occurs in tropical montane rain-
This species was named after the botanist and plant forest and high montane cloud forest up to the tree
892 collector Henry Hurd Rusby (1855–1940). line. In Peru it was found in lower montane rainfor-
est in Cusco, between 700 m and 1300 m a.s.l., and
near Machu Pichu in high montane forest at 2900
Vernacular names
m a.s.l. In Bolivia, where the species is much more
pino blanco (Bolivia) common and widespread, its altitudinal range is be-
tween 1500 m and 3350 m a.s.l. It is commonly as-
sociated in the forest with Meliaceae, Myrtaceae and
Description
other angiosperm trees; at high altitudes it forms
Trees to 25 m tall; trunk to 60 cm d.b.h. Bark on thickets with other dwarfed trees and shrubs. It oc-
larger trees becoming fissured and scaly, brown cupies similar habitats as P. parlatorei within the
weathering grey. Branches spreading or ascending, same altitudinal range, but in Bolivia the two species
forming a broad open crown in the forest, more are geographically separated.
bushy in open terrain at high altitudes. Foliage
branches slender or stout, terete, grooved between
Conservation
remote leaf bases, terminating in broad conical
buds, on stout branchlets 5–7 mm wide at base, with There is some evidence of forest decline in the re-
lanceolate-acuminate outer scales 5–10(–16) mm gion, but we cannot ascertain its impact upon this
long and spreading out, the inner scales smaller and species. The Conifer Specialist Group nevertheless
erect or converging, buds on slender lateral branch- wish to flag it as NT, because it appears to have a
lets smaller, with shorter scales. Leaves on saplings limited distribution within Bolivia, with an estimat-
and shaded branches (in forest) larger than on ma- ed extent of occurrence (EOO) of 22,500 km2 and
ture trees and exposed small trees, 4–9 cm long, one location in Peru at ca. 300 km distance. Two lo-
6–10 mm wide. Leaves on mature and exposed trees calities from where collections were made are within
(2–)3–5 cm long, (4–)5–8(–10) mm wide, patent, protected areas: Parque Nacional de Choquecamiri
elliptic to linear-lanceolate, widest in middle part, in Bolivia and Machu Pichu National Park in Peru.
gradually or sometimes more abrubtly narrowing IUCN: NT
to a short petiolate base; margins slightly revolute,
gradually tapering to an acute apex, coriaceous, deep
Uses
green above, pale green below. Midrib a continuous
narrow groove on adaxial (upper) side, raised but No commercial uses are recorded of this species. Its
narrow (0.4 mm wide) and continuous to apex on wood is used locally for construction purposes. This
abaxial side. Stomata small, in numerous intermit- species is not known to be in cultivation.
tent lines on abaxial side. Pollen cones not observed,
presumably solitary. Seed cones axillary, solitary on
Podocarpus salicifolius Klotzsch & H. Karst. ex
3–10 mm long peduncles; receptacles composed of
Endl., Syn. Conif.: 209. 1847. Type: Venezuela:
an axis with two unequal, spreading bracts, fusing
Cordillera de la Costa, Distrito Federal, Gran
and swelling to 6–8 mm long with exserted, trian-
Colombia, [“Habitat in Colombia”], H. Karsten s.n.
gular bract tips, green becoming red. Seeds single
(holotype W, destroyed, isotype LE). Pl. 38
between bract tips, ovoid-globose, including the
epimatium ca. 7 mm long with an obtuse distal crest, Podocarpus pittieri J. T. Buchholz & N. E. Gray, J. Ar-
dark green. Seed proper not observed. nold Arbor. 29: 130. 1948.
 Etymology Distribution
The species epithet compares the leaves with those Bolivia; NW Brazil; Colombia; Peru; N Venezuela.
of willows (Salix). TDWG codes: 82 VEN 83 BOL CLM PER 84 BZN

Vernacular names Ecology

pinabete (Venezuela) Podocarpus salicifolius occurs in the Andes ranges
in montane to upper montane rain forest or cloud
forest, at altitudes between 1200 m and 2150 m a.s.l.
Description
When growing in dense rainforest it can attain 30 m 893
Trees to 30 m tall (often much smaller); trunk to 80 as a canopy tree, but in cloud forest on steep slopes it
cm d.b.h. Bark thin, smooth in young trees, even- usually remains much smaller, up to 12 m tall.
tually becoming shallowly fissured and exfoliating
in small strips, brown, weathering grey; inner bark
Conservation
pinkish. Branches spreading or ascending, forming
a broad crown in large trees. Foliage branches stout, Although uncommon in most areas where it occurs,
terete, grooved between leaf bases; terminating in Podocarpus salicifolius has a wide range in the An-
broad-based, conical buds with spreading, long acu- dean Cordillera from Venezuela to Bolivia. Despite
minate scales, the outer scales to 12 mm long, up to its early description, this species seems to be under-
4 mm wide above their base, with scarious margins, collected, except in northern Venezuela, and it is
a few sometimes elongated to leaf-like appendices likely to be discovered in other localities than pres-
20–25 mm long, the inner scales usually shorter. ently known within its extent of occurrence (EOO).
Leaves on saplings spreading on vigorous shoots, to It is therefore not considered to be threatened at
23 cm long and 17 mm wide. Leaves on mature trees present.
usually drooping but sometimes spreading, (7–)8–13 IUCN: LC
cm long, 8–12 mm wide, straight or falcate (droop-
ing leaves curved above the short, twisted, petiolate
Uses
base), lanceolate-linear, gradually narrowing at both
ends; margins flat; apex acute; leaf texture lax; leaf No commercial uses are reported of this species, but
colour green above, paler green below. Midrib on large trees will undoubtedly be logged for their tim-
adaxial (upper) side obtusely raised, continuous or ber, as are other podocarps. The wood is suitable for
fading towards apex, raised at least near base on construction, general carpentry, and furniture mak-
abaxial side. Stomata very small, in numerous in- ing. It is not known to be in cultivation.
termittent lines on abaxial face of leaf. Pollen cones
axillary, sessile, solitary from buds with short acu-
minate scales, long cylindrical, 40–80 mm long, 3.5– Podocarpus salignus D. Don, in Lambert, Descr.
4.5 mm wide; microsporophylls broadly ovate with Pinus 2: [20]. 1824. Type: Chile: [“Gen. novum de
mucronate apex, bearing two globose pollen sacs. Chile”; (on reverse) “Chili. Herb: Pavon”], R. Ruiz
Seed cones axillary, solitary on 10–25(–30) mm long, & J. A. Pavón y Jiménez s.n. (holotype BM). Fig. 288
slender, pendulous peduncles; receptacles formed
of an axis with two unequal, fused bracts with free,
Etymology
curved bract tips, swelling to 8–12 mm long and 5–7
mm wide distally, ripening from dark green to dark The species epithet compares the species with wil-
purple. Seeds solitary on largest fertile bract, includ- lows (Salix).
ing the epimatium ca. 10 mm long, ovoid-globose
with a distal crest terminating in a small protrusion.
Vernacular names
Seed proper not observed.
mañio, manique (Chile)
 m a.s.l., with the highest localities in the Andes on
Description
­S-facing slopes.
Trees to 20 m tall; trunk monopodial or branching
low, up to 1 m d.b.h. Bark thin, smooth in young
Conservation
trees, becoming ragged with exfoliating strips on
older stems, light brown weathering whitish grey. In the northern part of its range there has been sub-
Branches in young trees forming a pyramidal crown, stantial decline due to conversion of natural for-
in larger trees spreading out more widely, forming a est to plantation forest with Eucalyptus and Pinus
broadly domed, often dense crown. Foliage branch- radiata. Increased fires have destoyed many other
lets slender, often drooping, terete, with fine grooves stands there. Cutting for firewood has also reduced
894 between decurrent leaf bases, terminating in small, the number of mature trees. In the south of its range
globose buds with imbricate, ovate-triangular scales. it is doing better, but there is a continuing decline of
Leaves on seedlings and saplings similar to those on its habitat resulting in reduced extent of occurrence
mature trees, narrowly linear-lanceolate, straight or (EOO) and area of occupancy (AOO) and, especial-
(slightly) falcate, (3–)5–9(–12) cm long, (3–)4–7 mm ly in the north, fragmented subpopulations. Habitat
wide, lax, narrowing to a short petiolate base, gradu- degradation is probably the most serious and wide-
ally tapering to an acute apex, lustrous light green spread menace to the survival of this species, which
above, pale green below. Midrib on adaxial (upper) does not react well to the opening up of forest cover
face less than 0.4 mm wide, raised from base but and concomitant dehydration in summer.
fading towards apex, similarly narrow but continu- IUCN: VU (A2 c-d)
ous to apex on abaxial side. Stomata very small, in
numerous intermittent lines on abaxial side. Pollen
Uses
cones axillary, sessile, solitary or grouped with two
or more, often from flushing leaves, or from older The fine-grained wood of this species is used locally
leaves, 25–50 mm long, very slender, the longest in the craft industry and was in the past of impor-
almost filiform, 1.5–2 mm wide at anthesis, yellow; tance for house building. More important now per-
microsporophylls minute, triangular, bearing two haps is its use in horticulture. Its date of introduction
globose pollen sacs. Seed cones axillary, solitary on in Europe is uncertain, but it has been very success-
slender, 1–2 cm long peduncles; receptacles formed ful and is quite common in large gardens and parks
of an axis with 2–3 fused bracts, one or two of these in regions with mild winters and abundant rainfall.
fertile, swelling to 5–6 × 4–5 mm, succulent deep It usually forms a bushy tree with dense, pendulous
red to violet. Seeds solitary or sometimes two on a foliage.
receptacle, including the epimatium 7–8 × 4–5 mm,
obliquely ovoid, green maturing to reddish, with an
Podocarpus salomoniensis Wasscher, Blumea 4 (3):
obtuse crest. Seed proper not observed.
430. 1941. Type: Solomon Islands: San Cristobal,
[“Hinuahaoro”], L. J. Brass 2881 (holotype B,
Distribution destroyed, isotypes A, BM, L).
S Chile: Biobio, La Araucania, Los Lagos, Maule.
Etymology
TDWG codes: 85 CLC-BI CLC-LA CLC-MA CLS-LL
The species epithet refers to the Solomon Islands,
where this species is endemic.
Ecology
In the north of its range (Mediterranean climate) Vernacular names
Podocarpus salignus follows water courses in the
dengali tolo (Malaita)
Roble-Hualo forest type (Bosque Maulino Costero);
in the Andes higher than 1000 m a.s.l. it can form
Description
pure stands in wetter areas away from water courses,
or it occurs in moist ravines. Here it is a minor com- Trees to 25 m tall, with a straight bole in forests to
ponent of Nothofagus obliqua forests. The altitudi- 35 cm d.b.h., stunted on poor growing sites; trunk at
nal range of this species is between 20 m and 1200 base not buttressed. Bark usually smooth, in larger
trees flaking with thin, curling strips, pale yellowish it may have disappeared. The other three subpopu-
brown; inner bark weakly fibrous. Branches spread- lations are represented by seven herbarium collec-
ing in tiers, sinuous, often drooping, forming a dif- tions from San Cristobal. Deforestation has mainly
fuse, conical crown, in old trees more irregular and affected the lower elevations on the islands of the
rounded. Foliage branches terete, stout, grooved archipelago. No recent collections have been made
between leaf bases, terminating in a large conical that came to the attention of the IUCN Conifer
bud with subulate-acuminate, keeled scales to 11 Specialist Group, the most recent (from San Jorge
mm long with spreading or curved tips. Leaves on Island) dates from 1965. This species is not known
saplings and young trees broader than on mature from any protected areas.
trees, to 20 mm wide, thin and flexible. Leaves of IUCN: EN [B2 a-b (iii)]
mature trees on distal parts of branches only, spread- 895
ing wide or more or less drooping, 10–25 cm long,
Uses
7–12 mm wide, linear-lanceolate, gradually tapering
to a narrow petiolate, twisted base (leaving crescent- This species, where it develops into a tree of moder-
shaped pulvini on the shoot when fallen) straight or ate size with a straight bole, is a useful timber tree
slightly falcate, tapering to an acute or obtuse apex, producing soft, light wood suitable for light con-
thick ­coriaceous. Midrib prominent on both sides of struction, interior finishing and boxes or cases. Lo-
leaf, extending the full length, more or less acute on cally it is being used to make oars. This species is not
adaxial (upper) side, 0.5–0.7 mm wide, obtuse, be- known to be in cultivation.
coming flattened distally and to 1 mm wide on abax-
ial side. Leaf colour lustrous green above, dull green
below. Stomata small and inconspicuous, in numer- Podocarpus sellowii Klotzsch ex Endl., Syn. Conif.:
ous intermittent lines on either side of abaxial mid- 209. 1847.
rib. Pollen cones unknown. Seed cones in upper leaf
axils on slender 10–15 mm long peduncles; recepta-
Etymology
cles subtended by 2 subulate, ca. 4 mm long, acute
bracts (‘foliola’), obconical, slightly flattened, 8–10 This species was named after the German botanist
mm long and broad, ca. 5 mm thick at apex, swelling Friedrich Sellow [Sello] (1789–1831).
to succulent red. Seeds 1, sometimes 2 per receptacle,
including the epimatium subelliptical, 11 × 8 mm,
Vernacular names
narrowed at base and rounded at apex, smooth and
glaucous green. Seed proper not observed. No common names have been recorded for this
species other than the generic ‘pinheirinho’, which
means little more than ‘conifer’.
Distribution
Solomon Islands, known from two islands [San
Description
Cristobal (Makira) and San Jorge Island near Santa
Isabel (Bughotu)]. Small trees to 10–12 m tall, d.b.h. to 30 cm. Branches
TDWG codes: 43 SOL-SO spreading, forming a rounded or irregular crown.
Foliage branchlets stout, terete, longitudinally
grooved between leaf bases, terminating in coni-
Ecology
cal buds 3–4 mm wide at base, with free, erect or
Tropical rain forest or swamp forest, the latter on spreading, apiculate outer scales which are some-
ultramafic soil and with codominant Casuarina sp. times much elongated (to 10–15 mm) and leaf-like.
Elevation from herbarium specimen labels 120–900 Leaves on saplings larger than on mature trees, 8–15
m a.s.l. cm long, 12–20 mm wide, straight or slightly fal-
cate, linear to lanceolate-linear. Leaves on mature
trees (2–)4–9(–11) cm long, (3–)7–11(–13) mm wide,
Conservation
narrowly elliptic, or lanceolate, lanceolate-linear to
This species is only known from two islands, one of more or less linear, straight or slightly falcate, gradu-
which (San Jorge) is a low-lying small island, where ally or more abruptly narrowing to a petiolate base,
 gradually tapering to an acute apex; margins only in shrubs and trees) grow with it in most locations and
the longest and narrowest leaves parallel for some epiphytes on trees are mostly mosses and lichens.
distance, flat or slightly revolute; texture coriaceous
or thin coriaceous; leaf colour more or less equal-
Uses
ly green on both sides. Midrib narrow (–0.5 mm),
continuous or sometimes fading towards apex, As a small tree this species has little commercial
raised in a shallow groove on adaxial (upper) side value for timber. It may locally be used for firewood
or forming a groove, more acutely raised from a flat or sometimes for fence posts. It is not known in cul-
leaf surface and continuous to apex on abaxial side. tivation.
Stomata small, in two bands of intermittent lines on
896 either side of abaxial midrib. Pollen cones axillary, 2 varieties are recognized:
solitary, sessile with obtuse perular scales near base,
narrowly cylindrical, 10–20 mm long, 1–2 mm wide; Podocarpus sellowii Klotzsch ex Endl. var. sellowii,
microsporophylls ovate-acuminate, with minutely Syn. Conif.: 209. 1847. Type: Brazil: [locality not
denticulate margins, bearing two globose pollen known], F. Sello s.n. [holotype B, destroyed, iso-
sacs. Seed cones axillary, solitary on slender, to 10 types F, GH, NY, UC).
mm long peduncles; receptacle small, swelling to
6–8 mm long, 4–6 mm wide, succulent reddish pur-
Description
ple turning dark brown. Seeds solitary, subglobose,
including the epimatium 7–9 × 5–8 mm, without a Leaves of mature trees larger than in var. angustifo­
crest. Seed proper globose with a smooth surface. lius, (3–)4–9(–11) cm long, (5–)7–11(–13) mm wide,
lanceolate to lanceolate-linear or nearly linear. Pol-
len cones ca. 20 mm long, 2 mm wide. Seeds 8–9 ×
Distribution
6.5–8 mm.
Brazil: Paraná, Rio Grande do Sul, Rio de Janeiro,
Santa Catarina, Sao Paulo.
Distribution
TDWG codes: 84 BZL-RJ BZL-SP BZS-PR BZS-RS
BZS-SC Brazil: Paraná, Rio Grande do Sul, Rio de Janeiro,
Santa Catarina, Sao Paulo.
TDWG codes: 84 BZL-RJ BZL-SP BZS-PR BZS-RS
Ecology
BZS-SC
Podocarpus sellowii has a coastal distribution in S
Brazil and occurs in the Atlantic Forest (Mata At-
Conservation
lantica). The remnants of this forest are situated in
the coastal mountains. These forests are evergreen Var. sellowii has a patchy occurrence in an area that
subtropical and grow on acidic soils derived from has a very high human footprint factor (GIS-AVG
granite and gneiss. Podocarpus sellowii is found in = 51) and where deforestation has been serious, in-
the montane forests at altitudes between 800 m dicating continuous decline. The extent of occur-
and 1800 m a.s.l. The forest type in which P. sellowii rence (EOO) calculated on the basis of herbarium
mostly occurs is relatively open and has an average specimen distributions is 188,608 km2 but there is a
canopy height of 10–15 m. In the northern half of its very scattered occurrence in coastal areas eastwards
range this occurs predominantly on eastern slopes, where the Atlantic Coastal Rain Forest has been
which receive less rainfall, and in the south on most decimated. Only one of 23 herbarium collections
slopes between 1100 m and 1500 m a.s.l. Podocarpus (from 16 localities) checked for the assessment has
sellowii may be associated with P. lambertii in the been recorded from a protected area, the Atlantic
south of its range, where gallery forest along rivers Forest Southeast Reserves.
becomes more prominent and intercalates with the IUCN: VU (A2, c)
Atlantic Forest. Palms and numerous dicots (both
Podocarpus sellowii Klotzsch ex Endl. var. angusti­
Description
folius Pilg., in Engler, Pflanzenr. IV.5 [18]: 88. 1903.
Type: Brazil: Rio de Janeiro, Nova Iguaçu, Pico do Trees to 30 m tall, with erect trunk to 80 cm d.b.h.
Tingua, A. F. M. Glaziou 17778 (lectotype K, here Bark thin, becoming scaly on large trees, slightly fi-
designated). brous, brown. Branches spreading to form a broad
crown. Foliage branchlets terete, smooth, finely
grooved, terminating in large, conical to fusiform
Description
buds 3.5–8 mm long and 2–4 mm wide at or above
Leaves of mature trees smaller than in var. sellowii, base, with imbricate, appressed, triangular or ovate,
(2–)3–5 cm long, 3–8 mm wide, narrowly elliptic to acute or obtuse and often mucronate scales. Leaves
lanceolate. Pollen cones ca. 10 mm long, 1 mm wide. on mature trees (3–)5–11(–13) cm long, (8–)10–18 897
Seeds 7–8 × 5–6 mm. (–20) mm wide, linear-lanceolate, usually straight
but sometimes slightly curved, tapering gradually
or in wider leaves more abrubtly to a short petiolate
Distribution
base; apex acute or sometimes obtuse; midrib ob-
Brazil: Rio de Janeiro (Cerro dos Orgaos, Pico do scurely raised on adaxial (upper) side, slightly less
Tingua). than 1 mm wide, more conspicuous on abaxial side;
TDWG codes: 84 BZL-RJ leaf colour lustrous dark green above, pale green
below. Stomata numerous, small, in two bands of
intermittent lines on abaxial side. Pollen cones axil-
Conservation
lary, solitary or with 2–3 together on 2–9 mm long
This rare variety is known from only two localities peduncles, cylindrical, 30–45 mm long, 4–6 mm
next to a metropolis. The locality Pico do Tingua wide; microsporophylls spreading, falcate, to 2 mm
is in a biological reserve (Reserva Biologica Tin- long with two basal globose pollen sacs. Seed cones
gua) and this variety occurs here frequently and has axillary, solitary on slender, 6–9 mm long pedun-
been regularly collected (specimens in herbarium cles; receptacles subtended by two very small bracts
RB at Rio de Janeiro seen by Dr Nicholas Hind of (‘foliola’), growing and swelling to 5–7 mm long and
the Royal Botanic Gardens, Kew, April 2008) since 5 mm thick at distal end, ripening to deep red, suc-
its discovery more than a century ago. The status of culent. Seeds solitary on distal part of receptacle, ob-
it in the other locality is unknown. Deforestation liquely attached, ovoid with a short proximal beak
has probably greatly reduced the area of occupancy and an acute distal crest, 12–15 × 8–10 mm when
(AOO) but the reserve seems to be well protected. mature; epimatium ripening from green to dark red.
IUCN: CR [B1 a-b (iii)] Seed proper not observed.

Distribution
Podocarpus smithii de Laub., Blumea 30 (2):
257. 1985. Type: Australia: Queensland, Atherton Australia: Queensland (Atherton Tableland, Mt.
District, Mt. Lewis, D. J. de Laubenfels P 464 Lewis).
(holotype A). TDWG codes: 50 QLD-QU

Etymology Ecology
This species was named after Lindsay Stuart Smith Podocarpus smithii is a rare species occurring in a
(1917–1970), plant collector in Queensland and New few localities in montane rainforest on the eastern
Guinea. Atherton Tableland and on Mt. Lewis at elevations
between 900 m and 1200 m a.s.l. It is most often
found growing along mountain creeks in acidic soils
Vernacular names
derived from granitic rock.
No common names have been recorded for this spe-
cies.
 Conservation Description
Podocarpus smithii was originally described from Small to medium sized trees 3–10(–20) m tall, up to
Mt. Lewis in northern Queensland and it was be- 30 cm d.b.h. Bark not described. Branches spread-
lieved to be restricted to that locality. Since 1985 ing, forming a rounded or domed crown. Foliage
other localities have become known in the Atherton branchlets terete, more or less grooved, glabrous,
Tableland and further north in Queensland, but it terminating in 2–6 mm long, 2–3 mm wide buds
is still considered to be a rare species occurring in with spreading, triangular outer scales. Leaves on
small and scattered populations. General deforesta- juvenile plants petiolate, elliptic to oblong, to 13 cm
tion in the area through logging and land use con- long and to 23 mm wide, straight, abrubtly narrowed
898 version of forested areas are likely to have reduced at base, acute or obtuse to rounded at apex. Leaves
its area of occupancy (AOO), but it remains difficult of mature trees more or less crowded at distal end of
to estimate to what extent this has taken place. This branchlets, spreading or ascending, elliptic to linear-
species should at least be flagged as Near Threatened oblong, 5–10 cm long, (8–)12–20 mm wide, straight
and is in need of a more thorough evaluation of its or slightly curved, petiolate at an abruptly narrowing
conservation status. base, with an obtuse or rounded (sometimes mucro-
IUCN: NT nate) apex, more or less coriaceous. Midrib obtusely
raised on adaxial side, ca. 1 mm wide, prominent and
wider (to 2 mm) on abaxial side; leaf colour dark
Uses
green above, pale green below. Stomata very small, in
Logging for timber would almost certainly have numerous irregular lines on abaxial (under) side on
included this species, as podocarp wood was and either side of midrib. Pollen cones axillary, in clus-
is considered valuable as construction material. At ters of 3, short pedunculate with several basal bract
present, native woodland in Queensland, and more scales, cylindrical (mature cones not observed). Seed
particularly rare species of trees in it, are considered cones axillary, solitary on a 2–6 mm long peduncle;
for protection and logging has therefore been sub- receptacles with 2 bracts (‘foliola’) 1–2 mm long at
stantially reduced. This species may occur in a few base, obliquely bilobate, 5 mm long, swelling with a
botanic gardens, but is otherwise not known to be truncate distal end, turning red and succulent when
in cultivation. ripe. Seeds including the epimatium ovoid, 7 × 5
mm, distal end obtuse, green turning purple or dark
brown when ripe. Seed proper not observed.
Podocarpus spathoides de Laub., Blumea 30 (2):
267. 1985. Type: Malaysia: Peninsular Malaysia,
Taxonomic notes
Johore, Gunung Ledang [Mt. Ophir], D. J. de
Laubenfels P 600 (holotype L). Podocarpus spathoides, first described and named
by De Laubenfels in 1985 (op. cit.), was based on his
Podocarpus spathoides de Laub. var. solomonensis collection P 600 from Gunung Ledang [Mt. Ophir]
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39. 2000. in Peninsular Malaysia. The isotype specimen at
K from a mature but small tree has elliptic to lin-
ear-oblong, obtuse or obtuse-mucronate to apically
Etymology
rounded leaves 5–8 cm long and 12–13 mm wide,
The species epithet comes from Latin spatha = a with a raised midrib on the upperside of the leaf
broad flat wooden or metal blade and refers to the and a flat to shallowly grooved midrib on the lower
shape of the leaves. (abaxial) side. On that mountain it occurs from the
upper limit of P. ridleyi (a species with lanceolate,
acute leaves) at ca. 1000 m to the summit at 1276
Vernacular names
m. It is a poorly known species based on very few
No common names are recorded for this species. ­collections.
 Distribution Vernacular names
Malesia: Peninsular Malaysia (Gunung Ledang = No common names have been recorded for this spe-
Mt. Ophir), Sarawak (Lawas), Maluku [Moluccas] cies.
(Morotai); Papuasia: Papua New Guinea (Rossel Is-
land), Solomon Islands.
Description
TDWG codes: 42 BOR-SR MLY-PM MOL 43 NWG-
PN SOL-SO Shrubs to 3 m tall, usually sprawling and not more
than 1.5 m tall, layering. Bark thin, fibrous, brown.
Foliage branchlets slender, terete, finely grooved, ter-
Ecology
minating in small, 2–4 mm long buds with narrowly 899
This species has been collected at widely disjunct lo- triangular scales ending in fine, spreading apices.
calities in Malesia, in various forested habitats. On Leaves on young and adult plants similar, narrowly
Gunung Ledang [Mt. Ophir] it occurs at altitudes linear-lanceolate to linear, 2.5–7 cm long, 2–4.5 mm
from ca. 1000 m to the summit at 1276 m a.s.l. in wide, straight or slightly curved, short petiolate or
low shrubby vegetation and stunted forest. It is ap- subsessile, gradually tapering to base and to an apicu-
parently rare there as on a visit in September 2008 late, pungent apex. Midrib acutely raised but narrow
with staff from FRIM I could not find it in the sum- (0.3 mm wide) on adaxial (upper) side, prominently
mit area. In the Solomon Islands it is found near sea raised and slightly wider on abaxial side. Leaf colour
level in evergreen tropical rainforest. yellowish green or lustrous green above, glaucous
green below. Stomata in two conspicuous bands on
either side of abaxial midrib, forming intermittent
Conservation
lines. Pollen cones axillary, solitary or in clusters of
The widely disjunct distribution is based on a very 2–5 on 1–4 mm long peduncles with minute scale
limited number of herbarium collections, either leaves, short cylindrical, 4–8 mm long, 1.5–2.5 mm
made by the author of this species, or identified as wide; microsporophylls minute, imbricate, finely
that species by him. More information based on apiculate, bearing two basal, globose pollen sacs.
more comprehensive collecting is needed to con- Seed cones axillary to reduced leaves, full-grown
firm its taxonomic status as well as to ascertain its leaves or subterminal, solitary on slender, 5–10 mm
distribution. It is unlikely to be limited to these few long peduncles; receptacles subtended by two nar-
localities, which span the length of the Malesian Ar- rowly lanceolate, curved or revolute 3–5 mm long
chipelago. bracts (‘foliola’), growing and swelling to 6–10 mm
IUCN: DD long, 6–7 mm wide, succulent and purple when ripe.
Seeds obliquely attached to distal end of receptacle,
ovoid when mature, 8–10 mm long, 5–7 mm wide,
Uses
often distally crested; epimatium glaucous, becom-
No uses have been recorded of this species. ing dark purple. Seed proper not observed.

Distribution
Podocarpus spinulosus (Sm.) R. Br. ex Mirb.,
Mém. Mus. Hist. Nat. 13: 75. 1825. Taxus spinulosa Australia: New South Wales, Queensland.
Sm., in Rees, Cyclopaedia 35, Taxus No. 7. 1819; TDWG codes: 50 NSW-NS QLD-QU
Margbensonia spinulosa (Sm.) A. V. Bobrov &
Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
Ecology
Biol. 103 (1): 60. 1998. Type: Australia: New South
Wales, Sydney Harbour, [“Port Jackson”], A. Philip Podocarpus spinulosus is a shrub growing on nutri-
s.n. (holotype LINN). Fig. 289, 290, 291 ent-deficient soils derived from sandstone and on
sandy old beaches and dunes along the coast as well
as farther inland on plateaux (Blackdown Tableland
Etymology
in New South Wales). On the coast it ­occurs in ‘heath
The species epithet refers to the spiny leaf tips. shrubland’ while in the interior it is found scattered
 in dry sclerophyll forest dominated by Angophora the outer scales enclosing the inner scales. Leaves
spp. and Casuarina spp. or with Eucalyptus gummi­ on saplings similar but slightly larger than on ma-
fera, E. sieberi, Acacia spp., Angophora bakeri, Dill­ ture plants, erect to patent, densele set on exposed
wynia retorta, Lambertia formosa, Leptospermum branches of older trees and more remote on shaded
attenuatum, etc., in sandy soil derived from sand- young trees, 2–4(–7) cm long, elliptical to linear or
stone, or on the rocks themselves. On the islands oblanceolate-linear, straight, 3–5 mm wide, gradu-
in Moreton Bay, Queensland, it grows on stabilized ally narrowing to a short petiolate or subsessile base,
dunes covered in shrubs and scattered eucalypts. gradually or more apruptly narrowing to an angus-
tate-acute or pungent (occasionally obtuse) apex;
margins slightly revolute; texture rigidly coriaceous;
Conservation
900 leaf colour lustrous dark green above, pale green
IUCN: LC below; flushing leaves glaucous green. Midrib on
adaxial (upper) side a continuous (shallow) groove,
sometimes raised proximally within this groove, on
Uses
abaxial side raised but fading towards apex. Stomata
This shrubby species of the genus Podocarpus ap- very small, in numerous intermittent lines on abax-
pears not to be in cultivation outside a few collec- ial face. Pollen cones in clusters of 3–8 or more due
tions in botanic gardens. Its ability to grow well in to corymbose branching at the end of 10–20 mm
poor sandy substrates near the sea and its layer- long, scaly or leafy stalks, sessile, subtended by short
ing habit would make it a useful stabilizer of drift- triangular bud scales, cylindrical, up to 10 mm long,
ing sand in warm temperate to subtropical climate. 2 mm diam.; microsporophylls broadly obtuse, with
Research into its potential for invasiveness in alien a scarious margin, bearing 2 small, globose pollen
countries and habitats should be conditional if in- sacs. Seed cones axillary, solitary on 3–7 mm long
troduction is considered in this context. peduncles; receptacles 6–7 mm long, consisting
of an axis with two unequal bracts, swelling only
slightly to become more or less succulent and pur-
Podocarpus sprucei Parl., in Candolle, Prodr. 16 plish red. Seeds globose, including the epimatium
(2): 510. 1868. Type: Ecuador: [“Valle de Pangor, 5–7 mm diam., with a minute, ridge-like crest.
ad 11,500’ ascendit”], R. Spruce 5519 (holotype not
located, isotypes BM, K). Fig. 292
Distribution
Equador, N Peru (Piura).
Etymology
TDWG codes: 83 ECU PER
This species was named after Richard Spruce (1817–
1893), who collected extensively in Ecuador and dis-
Ecology
tributed specimens to many herbaria.
Podocarpus sprucei is a high montane to subalpine
species growing in cloud forest up to the tree line, at
Vernacular names
altitudes from 1800 m to 3900 m a.s.l. Only in forests
Guabisay, Romerillo (Ecuador) at lower altitudes does it become a tree to 20 m tall.

Description Conservation
Trees to 20 m tall (dwarfed at highest altitudes); Overexploitation for timber, logging the larger
trunk to 50 cm d.b.h. Bark becoming scaly, reddish trees, was observed in the field by the plant collec-
brown. Branches ascending or spreading, forming a tor W. H. Camp (vicinity of Cuenca, Province of
bushy crown. Foliage branchlets slender, terete, fine- Azuay, Ecuador) in 1945, resulting in scarcity of the
ly grooved between leaf apices; terminating in clus- resource in that area. This species, last evaluated in
ters of small, verticillate buds or single, subglobose the 1990s and then considered not at risk, should be
buds with imbricate, broadly triangular, carinate reassessed using the 2001 IUCN criteria Version 3.1.
scales with denticulate margins and acuminate apex, IUCN: LC
 of abaxial midrib. Pollen cones axillary, solitary, ses-
Uses
sile, subtended by acuminate bud scales, cylindrical,
The wood of this species is much in demand and length at mature stage not observed. Seed cones ax-
trees are logged from primary forest often unsus- illary, solitary on 7–18 mm long peduncles; recep-
tainably due to scarcity and slow growth. It is used tacles composed of an axis with two unequal, fused
in house construction and to make furniture as it and slightly swollen bracts, 7–8 mm long, becoming
takes a fine polish. This species is also planted as an red or purple. Seeds including the epimatium ovoid-
ornamental in parks in Ecuador. globose, ca. 8 mm long, with a minute crest.

Podocarpus steyermarkii J. T. Buchholz & Distribution

901
N. E. Gray, J. Arnold Arbor. 29: 133. 1948. Type:
Guyana: Pakaraima Mts.; Venezuela: Bolivar (Car-
Venezuela: Bolívar, Rio Carrao, Carrao-tepui,
rao-tepui, Uaipan-tepui, Cerro Jaua), Amazonas
J. A. Steyermark 60876 (holotype F). Pl. 39
(Neblina Massif).
TDWG codes: 82 GUY VEN
Etymology
This species was named after Julian A. Steyermark
Ecology
(1909–1988) of the Field Museum of Natural History
in Chicago, who hugely contributed to the botany of Podocarpus steyermarkii is a rare species occurring
Central and South America. on the upper slopes and summits of the tepuis (table
mountains) of Guyana and Venezuela. Its altitudinal
range, as known from as yet limited collections in
Vernacular names
herbaria, is from 1850 m to 2430 m a.s.l. This species
No common names have been recorded for this spe- has now been found in cloud forest with a canopy to
cies. 20–25 m tall, as well as in low herbaceous or scrub
vegetation in areas with blanket bogs with grasses,
ferns, pitcher plants, orchids, mosses, and low scat-
Description
tered shrubs to 1.5 m tall, interspersed with thickets
Small trees to 12 m tall, usually stunted in habit; and dwarf forest including palms. Here the podo-
trunk to 30 cm d.b.h. Bark undescribed, probably carps (P. roraimae and P. tepuiensis also occur in this
thin, smooth, becoming somewhat scaly on larger habitat) occur mostly in the dwarf forest patches
trunks, brown. Crown densely branched. Foliage along small streams, which are the sources of large
branches stout, rigid, terete, with grooves running river systems. The substrate is acidic peat or peaty
down from slightly raised leaf bases, terminating forest soil over sandstone. Levels of precipitation
in large buds 4–7 mm wide at base, with broadly are very high and there is frequent, long lasting fog
ovate, scariously margined outer scales terminating (clouds surrounding the tepuis), causing low tem-
in a 5–15 mm long, cuspidate and erect or spread- peratures in the daytime.
ing apex (sometimes one or more scales becoming
even longer and leaf-like); inner scales shorter with
Conservation
converging apices. Leaves more or less remotely in-
serted, patent, (3–)4–8 cm long, straight or slightly This species, which was described and known from
falcate, (6–)8–11 mm wide, thick coriaceous, lanceo- only a single locality in 1948, is still known from
late to lanceolate-linear, narrowing to a petiolate, only six or seven localities. It is likely to turn up in
often twisted base, slightly V-shaped transversely more places with continuing botanical surveys, but
or nearly flat; margins slightly revolute, gradually is obviously scattered and uncommon. Its total area
tapering to an acute or obtuse apex; leaf colour lus- of occupancy (AOO) is almost certainly less than
trous dark green above, paler green below. Midrib 500 km2, but it is not exploited, nor are the forest
on adaxial (upper) side slightly raised with a central patches in which it occurs threatened by deforesta-
groove, or only a groove continuous to apex, obtuse- tion. This species is therefore considered to be out-
ly raised and 1 mm wide on abaxial side. Stomata side any category of threat of extinction in the wild.
small, in numerous intermittent lines on either side IUCN: LC
902

6 4

7
3

Pl ate 39 . Podocarpus steyermarkii. 1. Branch with leaves and seed cones. 2. Seed cone. 3. Branch with
leaves and pollen cones. 4. Microsporophyll (dried). 5. Foliage bud and leaves. 6. Bud scale. 7. Cross-section
of leaf.
 the green epimatium, minutely crested distally, rip-
Uses
ening to blackish purple. Seed proper ca. 8 × 7 mm,
No uses have been recorded of this species and it is slightly flattened, brown.
not known to be in cultivation.
Taxonomic notes
Podocarpus subtropicalis de Laub., Blumea 30 (2): Podocarpus subtropicalis has been identified with
277. 1985. Type: China: Sichuan, Emei Shan, [“Mt. P. neriifolius by Chinese botanists (see Flora of Chi-
Omei”], E. H. Wilson 3007 (holotype A). na 4: 82, 1999), although it is usually not explicitly
synonymized with that species in the recent floristic
Podocarpus subtropicalis de Laub. var. medogensis literature. Its taxonomic status remains somewhat 903
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 40. 2000. uncertain and is not helped by the circumstance that
it seems to have been widely spread in cultivation.
Its morphology is also close to P. fasciculus, a species
Etymology
from Taiwan described by De Laubenfels (1985) in
The species epithet refers to the subtropical habitat the same paper on the same page, both with short
or distribution of this species. Latin descriptions and type designations, but with-
out comparative discussion. I have retained these
species here as distinct, but more critical taxonomic
Vernacular names
comparison based on further sampling from trees
No vernacular name is recorded for this species. occurring in the wild is needed.

Description Distribution
Trees to 20 m tall, monopodial. Bark not described China: Sichuan (Emei Shan [Mt. Omei]), Yunnan.
or observed. Foliage branches terete, grooved, ter- TDWG codes: 36 CHC-SC CHC-YN
minating in buds of 4–5(–7?) × 2–3 mm, with lan-
ceolate, keeled and spreading or recurved scales.
Ecology
Leaves linear-lanceolate, petiolate (petioles 3–7 mm
long), on juvenile plants up to 18 cm long and 14 mm This species was described and named in 1985 by
wide; on adult plants 5–12(–15) cm long, (5–)7–13 David J. de Laubenfels, based on a collection from
mm wide, coriaceous, stiff, straight or more or less the famous mountain with Buddhist shrines and
falcate, gradually tapering at both ends, acute or stairway paths Emei Shan [Mt. Omei] in Sichuan,
acuminate. Midrib prominent on both sides, acutely China. Little is known of the ecological circum-
raised adaxially, wider and centrally grooved abaxi- stances and many trees have been introduced there
ally. Leaf colour bright green above (adaxially), pale over the centuries. It was subsequently reported
green below; new leaves flushing yellowish green. from Yunnan, but it remains a poorly known taxon.
Stomata on abaxial (under) side, very small, in nu- It is also widely planted elsewhere in China and, ac-
merous irregular lines on either side of midrib. Pollen cording to David de Laubenfels, in many other parts
cones axillary, sessile, in clusters of (1–)2–5, subtend- of the world with a warm-temperate to subtropical
ed by scarious, rostrate, keeled scales 3–5 mm long, climate.
cylindrical; cones elongating to 20–40(–60?) mm
× 2.5–3.5 mm at anthesis; micro­sporophylls spirally
Conservation
arranged, with a triangular, spreading, 0.3 mm long
apex and with two pollen sacs. Seed cones solitary IUCN: DD
and axillary on 5–15 mm long, slender peduncles;
receptacles 7–12 mm long, with two 1–2 mm long
Uses
bracts (‘foliola’) at their base, swelling when ripe to
a succulent, red pseudo-fruit ca. 10 mm thick. Seeds This species, occurring on Emei Shan [Mt. Omei],
single (sometimes 2) at truncate distal end of recep- has been taken from there and seeded and/or planted
tacle, obliquely globose, 10–12 × 8–10 mm including
 widely in other parts of China and beyond. It is often bud scales, cylindrical, 10–20 mm long, 3 mm wide,
identified as P. neriifolius. yellow turning brown; microsporophylls imbricate,
peltate, with erose-denticulate margin and minutely
apiculate, bearing two basal, globose pollen sacs.
Podocarpus sylvestris J. T. Buchholz, Bull. Mus. Seed cones axillary and solitary on slender, 7–15 mm
Hist. Nat. (Paris), sér. 2, 21: 285. 1949. Type: New long peduncles; receptacle subtended by two small
Caledonia: Grande Terre, Province Sud, Forêt du bracts (‘foliola’), growing to 6–10 mm long, swell-
Mois de Mai, J. T. Buchholz 1696 (holotype ILL). ing to 5–6 mm thick, maturing to red or red-brown.
Seeds including the epimatium relatively large,
Podocarpus novae-caledoniae Vieill. var. colliculatus 10–15 × 7–9 mm, obliquely ovoid with a faint crest,
904 N. E. Gray, J. Arnold Arbor. 39: 432. 1958; Podocarpus smooth, olive green turning brown. Seed proper
colliculatus (N. E. Gray) de Laub., Adansonia, ser. 3, ovoid, smooth, ca. 9 × 6 mm.
27 (2): 152. 2005.
Taxonomic notes
Etymology
De Laubenfels (op. cit.) elevated var. colliculatus of
The species epithet sylvestris means ‘from the forest’ N. E. Gray (op. cit.), which she had placed with Podo­
and refers to its habitat. carpus novae-caledoniae and described from the Île
des Pins in New Caledonia, to a species. Gray should
have placed this taxon (as a variety) with P. sylves­
Vernacular names
tris, from which it differs only slightly in having
No common names have been recorded of this spe- somewhat larger receptacles and longer peduncles.
cies. Both P. novae-caledoniae and P. sylvestris occur on
the Île des Pins and are well distinguished morpho-
logically, but the differences between P. colliculatus
Description
and P. sylvestris are minor and not constant in the
Small to medium size trees to 20 m, rarely to 30 m specimens I have seen.
tall and 80 cm d.b.h., or at high altitude shrubby
trees 1.5–3 m tall. Bark on large trees thick and fi-
Distribution
brous, fissured longitudinally, or on small trees
thin and more or less tesselated, brown. Branches New Caledonia (mountainous parts of Grande
on trees spreading to form a broad crown, densely Terre, Île des Pins).
set on shrubby trees, spreading out wide. Foliage TDWG codes: 60 NWC
branchlets terete, finely grooved, terminating in
small, globose or ovoid buds with 1–2 mm long,
Ecology
imbricate, appressed and obtuse scales. Leaves on
juvenile plants larger than on mature plants, to 16 Podocarpus sylvestris grows as a scattered tree in
cm long and 18 mm wide. Leaves on mature trees the tropical evergreen rainforests that cover parts
and shrubs variable, (3–)5–10(–13) cm long, (5–)7– of the mountains of New Caledonia; it is also found
11(–13) mm wide, oval-linear (widest at midpoint) in wooded ravines and at high altitude in vegetation
to linear, tapering gradually down to a more or less bordering on ‘maquis minier’, the shrubby vegeta-
petiolate base and to an obtuse apex, coriaceous, tion on serpentine. It is not restricted to these ultra-
straight or only slightly curved. Midrib prominent mafic soils, as it also occurs on micaschist in the far
but obtusely raised on adaxial (upper) side, 1–1.2 NE of the main island Grande Terre. The altitudinal
mm wide, flat or forming a groove in dried leaves range is 150 m to 1200 m a.s.l. At high altitude or on
on abaxial side. Leaf colour bright lustrous green ultramafic rocks it becomes a shrub or shrubby tree
above, dull light green below. Stomata very small, of low stature; in well developed rain forest it can
in two bands forming numerous intermittent lines reach 20–30 m tall. Besides numerous angiosperms,
on abaxial side. Pollen cones from clustered buds in its associated species are sometimes conifers, main-
axils of leaves or below these, sessile, solitary or 2–3 ly Araucaria spp. and Agathis spp. It is most often
together, subtended by small, rounded, imbricate found as a small understory tree with these conifers,
but is fertile in these circumstances just as in more leaf bases, terminating in small, globose buds 2–3
open maquis-like vegetation or as a canopy tree. mm long and wide, with inbricate, broadly trian-
gular, carinate scales with narrow scarious margins.
Leaves densely set, spreading at wide angles to shoot;
Conservation
on juvenile plants up to 6 cm long and 8 mm wide;
This species has been the target of logging, an ac- on mature plants (1–)1.5–2.5(–3.5) cm long, oblan-
tivity which has undoubtedly led to a reduction in ceolate or elliptic, 3.5–5 mm wide, often slightly or
its abundance, but is difficult to quantify. Such op- markedly curved down, gradually narrowing to a
erations have now virtually ceased. This species is short petiolate base; margins slightly revolute; apex
widely distributed and still locally common and if obtuse or more or less acute; leaf texture coriaceous;
forests are allowed to regenerate it will again spread leaf colour deep lustrous green above, pale green 905
from these localities. below; young leaves flushing pinkish white. Midrib
IUCN: LC on adaxial (upper) side of leaf a continuous groove,
inconspicuous and nearly flat on abaxial side. Sto-
mata very small, in numerous intermittent lines on
Uses
abaxial face of leaf. Pollen cones axillary, solitary,
This species has been subject to logging in the past; sessile with rounded perular scales at base, cylindri-
its wood provides good material for construction cal, 10–15 mm long, 1.5 mm wide, cream coloured
and indoor carpentry, floor boards, and furniture when mature; microsporophylls triangular, with
and has been known as ‘false kauri’ in the timber scarious margins, bearing two globose pollen sacs
trade. Exploitation has virtually ceased as the more at base. Seed cones axillary, solitary on a short (2–3
accessible forests have been logged and forest con- mm long) peduncle; receptacles constituting of an
servation of the remainder is now the prevailing axis with two unequal, divergent but mostly fused,
management practice. This species has been planted slightly swollen, fleshy bracts, the largest fertile, ca.
in some parks and gardens in New Caledonia. 5 mm long, green pruinose ripening to red or pur-
ple. Seeds including the epimatium 6–7 mm long, 4
mm wide, obliquely ovoid, with a distal crest form-
Podocarpus tepuiensis J. T. Buchholz & ing a papillate protrusion.
N. E. Gray, J. Arnold Arbor. 29: 134. 1948. Type:
Venezuela: Bolivar, La Gran Sabana, Ptari-tepui,
Distribution
J. A. Steyermark 59716 (holotype F).
Ecuador: Cordillera del Condor; Venezuela: Bolivar,
Amazonas.
Etymology
TDWG codes: 82 VEN 83 ECU
The species epithet refers to the (region of the) tepu-
is or table mountains of Venezuela.
Ecology
Podocarpus tepuiensis occurs in the mountains
Vernacular names
(tepuis = table mountains and other mountains) as
No common names have been recorded for this spe- well as in the lowlands. The altitudinal range in the
cies. mountains is from 700 m to 2450 m a.s.l.; the two
herbarium collections in the southern Venezuelan
lowlands are from 100–110 m a.s.l. In the latter lo-
Description
cality it occurs on the edge of forest near savanna,
Shrubs or small trees 1–6 m, rarely to 15 m tall; on white sand with seasonal flooding (black water),
trunks on the largest trees to 30 cm d.b.h. Bark thin, and with occasional fires on the savanna; associ-
fibrous, reddish brown. Branches ascending, form- ated species are e.g. Henriquezia nitida, Leopoldina
ing a (narrowly) pyramidal, dense crown in small ­piassaba, Emmotum sp., and Aldina sp. On the Ven-
trees, spreading more horizontally in shaded trees ezuelan tepuis it occurs on slopes and summits,
and with more or less erect foliage in shrubs. Foliage usually along streams in gallery forest or at higher al-
branchlets slender, terete, with fine grooves between titudes in dwarf forest patches. It may sometimes be
 a­ ssociated with P. roraimae and P. steyermarkii, but fibrous, orange-brown. Branches spreading, form-
these two podocarps do not occur at lower altitudes ing a rounded crown. Foliage branchlets terete,
than 1800 m a.s.l. and in most localities only one of slightly grooved or ridged, terminating in globose
these species has been found. The disjunct locality compact buds of 3–6 mm diam. with imbricate,
in Ecuador, where this species has been found since rounded scales with erose margins. Leaves on ju-
1993 (specimens identified by David de Laubenfels), venile plants lanceolate, to 20 cm long and 22 mm
is at an altitude of around 2000 m, but no specifica- wide, straight or falcate, thin, gradually tapering to
tion of the habitat or vegetation was given with the an acute apex. Leaves on mature plants lanceolate
collected specimens. to linear-lanceolate, 7–14 cm long, (10–)12–17(–20)
mm wide, straight or slightle curved, thinly coria-
906 ceous, gradually tapering to a petiolate base; apex
Conservation
acute or acuminate. Midrib on adaxial side nar-
Podocarpus tepuiensis is the most common and row, acutely raised from base to apex, on abaxial
widespread of the ‘tepuis podocarps’ and actually side wider, to 1.5 mm, obtusely raised. Stomata very
occurs elsewhere, too. It is a shrub or small tree of small, in numerous irregular lines on abaxial (un-
little interest for timber and most of the populations der) side. Pollen cones axillary, solitary or in clus-
are remote from roads and towns. The disjunct oc- ters of 2–3, sessile, subtended by rounded bracts
currence in Ecuador makes it likely that its total (bud scales), narrowly cylindrical, 25–40 mm long,
range is incompletely known at present. ca. 3 mm wide; microsporophylls spirally arranged,
IUCN: LC spreading, with triangular apex and two basal pol-
len sacs. Seed cones axillary, solitary on a 3–10 mm
long, slender peduncle; receptacles subtended by 2
Uses
minute, early deciduous bracts (‘foliola’), slender,
No uses have been recorded of this species and it is ca. 10 mm long, consisting of two lobes (axes?) each
unknown in cultivation. with a distal bract, one of these being fertile; lobes
swelling mostly distally, ripening to purplish pru-
inose. Seeds single, obliquely attached, including the
Podocarpus teysmannii Miq., Fl. Ned. Ind. [Fl. Ind. epimatium 7–9 × 5–7 mm, subglobose with a small
Batavae] 2 (7): 1072. 1859. Podocarpus neriifolius crest, dark green turning blackish brown.
D. Don var. teysmannii (Miq.) Wasscher, Blumea
4 (3): 453. 1941. Type: Indonesia: Sumatera, Utara,
Distribution
on the coast near Sibolga, J. E. Teijsmann s.n. (holo-
type L). Malesia: Peninsular Malaysia, Sumatera (including
Bangka & Lingga Islands), Borneo (Brunei, Sabah).
TDWG codes: 42 BOR-BR BOR-SB MLY-PM SUM
Etymology
This species epithet commemorates the Dutch bota-
Ecology
nist Johannes Elias Teijsmann (1809–1882).
In Flora Malesiana, ser. 1, 10 (3): 407 (1988) Podo­
carpus teysmannii was described as an under­storey
Vernacular names
tree to 12 m tall, but in fact sometimes trees are
On the west coast of Sumatera this species is known found reaching the canopy with a height of 36 m
as kalek rotan; Pilger (1903) cited the name Sikuju (T. C. Whitmore FRI 8835 collected on Gunung Blu-
laut, presumably also from Sumatera. mut in Peninsular Malaysia at 510 m on a ridge).
This species occurs in both primary and secondary
rain forest from near sea level to occasionally 1350 m
Description
a.s.l., but mostly up to 800 m. It is often restricted to
Trees to 36 m tall, commonly to 20 m, d.b.h. to 50 sites with poor, sandy, or rocky soils and therefore
cm; trunk erect, straight, terete; large trees with will not attain large size under these conditions. On
slightly fluted base. Bark smooth, eventually with the coast of Sabah it often occurs on sandy ridges or
small, powdery scales, brown; inner bark soft and hills with the broadleaved tree Dryobalanops rappa
(Dipterocarpaceae); in the Mt. Kinabalu area (Mesi- of large trees), straight or slightly curved, tapering
lau River) it was collected in ‘kerangas’ forest domi- to a slightly twisted base and an acute-pungent or
nated by Agathis sp. sometimes obtuse-mucronate apex, coriaceous, stiff;
a faint groove through the middle adaxially (upper-
side) and an obtuse midrib abaxially; leaf colour
Conservation
dark green, more or less lustrous above, dull and
IUCN: LC lighter green below. Stomata in two irregular bands
of intermittent lines on abaxial side. Pollen cones
axillary, solitary or sometimes in pairs on short
Uses
peduncles, subtended by small, triangular scales,
Larger trees of this species are undoubtedly logged cylindrical, 12–15(–20) mm long, 3–4 mm wide at 907
for their timber; the use of which will be similar to anthesis, straight or curved after shedding pollen,
that of other ‘podocarps’ of the region. Podocarp yellowish green turning brown; microsporophylls
wood is suitable for light construction, carpentry imbricate, peltate-triangular, upper margin denticu-
and joinery, furniture making, etc. late, with two sublateral, globose, yellow pollen sacs.
Seed cones axillary, solitary on a short peduncle;
receptacles when growing narrowly oblong, with
Podocarpus totara G. Benn. ex D. Don, in Lambert, two distal bracts subtending 1–2 ovules, swelling to
Descr. Pinus, ed. 2: 184. 1828. [& in ed. 8°, 2: [189]. nearly globose, 4.5–5.5 mm long, yellow ripening to
1832]. Type: New Zealand: North Island, Bay of scarlet, soft and succulent. Seeds including the epi-
Islands, [?] Bennett s.n. (lectotype BM, [specimen matium ovoid-oblong, with a constricted distal part,
a] designated here). Fig. 293, 294, 295 3–3.5 × 2–2.5 mm, lustrous green or glaucous green.
Seed proper ca. 2.5 mm long, narrowly ovoid; seed
Podocarpus totara G. Benn. ex D. Don var. waihoen­ coat smooth and hard.
sis Wardle, New Zealand J. Bot. 10 (1): 201. 1972.
Taxonomic notes
Etymology
A herbarium sheet of this species at BM contains a
The species epithet uses the Maori name for this tree. mixture of specimens collected by different people
at different localities and times. The largest branch-
let marked a) was collected by Bennet in July 1829 at
Vernacular names
the Bay of Islands. Brian Molloy annotated this as
totara (Maori) the lectotype of Podocarpus totara in October 1991,
but since he never published this, that lectotypifica-
tion is effected here.
Description
Trees to 30(–40) m tall, with a straight, some-
Distribution
times massive trunk to 3.6 m d.b.h. Bark thick,
fibrous, stringy, with longitudinal furrows, exfolia- New Zealand: North Island, South Island.
ting in small, fragmentary strips, light grey-brown. TDWG codes: 51 NZN NZS
Branches spreading, forming a rounded crown, in
veteran trees irregular through much reiteration.
Ecology
Foliage branchlets terete, glabrous, finely grooved;
branching irregularly, sometimes dense, but shaded Podocarpus totara was the largest conifer dominat-
branchlets more or less pectinate. Terminal buds 2–3 ing the lowland forests that covered much of the two
mm long, subglobose, the inner scales imbricate and large islands of New Zealand before humans, and es-
obtuse, the outer scales acute with free tips. Leaves pecially Europeans, intervened. It now occurs only
sessile, short decurrent, spreading at wide angles, in forest fragments, often in secondary vegetation. It
lanceolate-linear, (0.5)1–2.5(–3) cm long, (1.5–)2– is a slow growing, emergent tree dependent in that
3.5(–4.5) mm wide (narrowest leaves on saplings ecosystem upon episodal events of disturbance ena-
and young trees, widest leaves on fertile branches bling massive regeneration. Past ­forest clearance, if it
 took place in what are now forest rerserves, can pro- the ­numerous sawmills that had sprung up every-
vide these conditions. Maximum age is ca. 800–1000 where, the remainder were wasted (Ogden & Stew-
years and one tree is estimated to exceed 1800 years. art in Enright & Hill, 1995). Timber export between
In primary forest this species is often associated 1850–1900 included totara, but its main source was
with other conifers, e.g. Dacrycarpus dacrydioides, kauri (Agathis australis) until that resource was al-
Dacrydium cupressinum, Prumnopitys ferruginea, most exhausted. The wood is reddish, but variable
Phyllocladus trichomanoides, and in Northland in hue, dense and straight-grained, and was used for
­Agathis australis. Angiosperm trees can also be com- general construction, joinery, furniture and cabinet
mon and are often varied. Usually Podocarpus totara making, as well as the coarser applications like rail-
grows well below the altitude where Nothofagus be- way sleepers, posts for communication lines, and
908 gins to dominate, but occasionally it penetrates into fencing. At present, native trees are virtually univer-
this zone at around 500 m a.s.l. Above this altitude sally protected in New Zealand and the exploitation
it is replaced by the similar species P. cunninghamii. of the wood of totara is much restricted, essentially
to dead trees outside reserves, and put to specialist
wood work only. In horticulture it remains uncom-
Conservation
mon, mostly restricted to arboreta and botanic gar-
Under the A criterion of IUCN’s Red List of Threat- dens. Although eventually growing to a large tree if
ened Species™, this species would have to be listed in left alone, it can be planted and trimmed for hedges.
a threatened category due to relatively recent major
decline, caused by logging and forest clearance in
the past 150 years. In New Zealand, this criterion, Podocarpus transiens (Pilg.) de Laub. ex Silba,
if applied to native trees, is not accepted by conser- Phytologia Mem. 7: 68 (1984). Podocarpus lambertii
vationists, due to more recent legislation that gives Klotzsch ex Endl. var. transiens Pilg., in Engler,
protection to most of the remaining forests and trees, Pflanzenr. IV.5 [18]: 86. 1903. Types: S Brazil, F. Sello
both within reserves and through legal bans on cut- s.n.; A. F. M. Glaziou 16335 (syntypes, not located).
ting and other means of clearance of native forest.
This policy has in fact already lead to an increase in Podocarpus transiens (Pilg.) de Laub. ex Silba var.
abundance (from an all-time low just before forest harleyi Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39.
clearance was prohibited), due to regeneration and 2000.
spread from seed trees into secondary vegetation
and pioneer phases of native forest. As demand for
Etymology
further conversion of forested land for agriculture
(diary, sheep farming) is presently waning, for the The species epithet transiens is Latin for ‘passing
foreseeable future Podocarpus totara and other low- over into’ and presumably refers to similarities with
land conifers are indeed out of danger. However, this P. lambertii.
situation is policy dependent and there is no abso-
lute guarantee that this policy will never change in
Vernacular names
the other direction in the future. The IUCN Red List
attempts to assess the conservation status independ- No common names have been recorded for this
ent of current or future (national) policies. species other than the generic ‘pinheirinho’, which
IUCN: LC means little more than ‘conifer’.

Uses Description
The Maori have used ‘totara’ (not exclusively P. to­ Small trees to 8–12 m tall, d.b.h. to 30 cm. Branches
tara) for many purposes, but especially construc- spreading, forming a rounded or irregular crown.
tion of houses. Europeans began large scale logging Foliage branchlets slender, terete, longitudinally
and forest clearing in the 19th century and much of grooved between leaf bases, terminating in small,
the wood of totara went into fence posts on the es- subglobose buds with imbricate, broadly triangu-
tablishing farms and later in that century for rail- lar, carinate, short apiculate scales. Leaves on sap-
way sleepers. Only the ‘best’ trees were sawn in lings similar but slightly larger than on mature trees.
Leaves on mature trees crowded, spreading at wide
Ecology
angles or more or less directed forward, (1.5–)2.5–
5(–6.5) cm long, (3.5–)4–7(–9) mm wide, narrowly This species is an uncommon component of mon-
lanceolate to oblanceolate, straight, gradually nar- tane forest remnants surrounded by ‘campo rupestre’
rowing to a short petiolate base, gradually or more (rocky grassland), a dryland vegetation in the inte-
abruptly tapering to an acute or obtuse apex; mar- rior. Nearer the Atlantic coast and in the south of its
gins not parallel for most of leaf length, straight range it has been found in remnants of Atlantic Rain
or slightly revolute; leaf texture coriaceous or thin Forest. The altitudinal range of this species, based on
coriaceous; leaf colour green or grey-green above, data on herbarium specimen labels, is 1000–1780 m
whitish green below. Midrib 0.5–0.7 mm wide, con- a.s.l. It may be associated with Podocarpus lambertii,
tinuous or sometimes fading towards apex, mostly especially in the southern parts of its range, but un- 909
forming a groove on adaxial (upper) side, more or equivocal evidence of this has not been recorded on
less raised from a flat leaf surface and continuous herbarium labels or in the literature, probably due
to apex on abaxial side. Stomata small, in numer- to identification problems, as most specimens were
ous intermittent lines forming two bands on abaxial identified as P. lambertii.
side. Pollen cones axillary, on short shoots or slen-
der peduncles up to 15 mm long in 1-several clus-
Conservation
ters of 2–6 or more, subtended by apiculate perular
scales, cylindrical, 6–12 mm long, 1.5–2 mm wide; This species appears to be quite rare and was pre-
microsporo­phylls ovate-triangular, with minutely viously considered to be endangered based on pre-
denticulate margins, bearing two globose pollen 1994 IUCN criteria. Due to very small numbers of
sacs. Seed cones axillary, solitary on slender, 5–10 trees in some of these locations, the area of occu-
mm long peduncles; receptacles small, swelling pancy (AOO) must be much smaller than 2000 km2,
to 6–7 mm long, 4–5 mm wide, succulent, purple, while populations appear always to be small in the
eventually chestnut brown. Seeds solitary, globose, ca. 10 locations recorded in the region, of which the
including the epimatium 4–5 × 4–5 mm, glossy most commonly visited is the Morro do Chapeu in
without a crest. Seed proper globose with a smooth Bahia. In several locations degradation of habitat
surface. and tree cutting for firewood are causing decline of
this species. Deforestation caused by cutting, fire
and grazing of domestic animals is the main threat.
Taxonomic notes
The tree cutting is mostly for firewood, much less
This taxon was originally described by Pilger (op. for timber.
cit.) as a variety of Podocarpus lambertii, but it was IUCN: VU [B2 a (iii, v)]
noted by David de Laubenfels to be sufficiently and
consistently different to be recognized as a species,
Uses
whereupon Silba (op. cit.) published the new combi-
nation and status. The main or obvious differences No commercial uses have been recorded for this
are in the leaf shape and size, but another difference small tree, of which the timber has not much value.
are the persistent, apiculate bud (perular) scales at It is used locally for fence posts and more extensive-
the base of the pollen cones. It occurs further north ly for firewood. It is not known to be in cultivation.
than P. lambertii, but there is one herbarium record
in K from Santa Catarina in the south, where P. lam­
bertii is most common. Podocarpus trinitensis J. T. Buchholz & N. E. Gray,
J. Arnold Arbor. 29: 135. 1948. Type: Trinidad and
Tobago: Trinidad, Saint Andrew Co., Valencia,
Distribution
[“Valencia road”], W. E. Broadway 7151 (holotype
Brazil (Bahia, Goias, Minas Gerais, Paraná, Santa MO).
Catarina).
TDWG codes: 84 BZC-GO BZE-BA BZL-MG BZS-PR
Etymology
BZS-SC
The species epithet refers to the island of Trinidad.
 narrows more abruptly and is often weakly apicu-
Vernacular names
late. Whether the pollen cones can reach the length
No common names have been recorded for this spe- found in P. coriaceus is not known, because only im-
cies. mature cones, not much beyond the bud stage, have
so far been observed in collected specimens. Podo­
carpus trinitensis may turn out to be only a short-
Description
leaved variety of P. coriaceus; both taxa are present
Trees to 15–20 m tall; trunk with d.b.h. to 60–80 cm, on the island of Trinidad, but P. coriaceus extends
often fluted at base. Bark smooth, on large trunks along the Lesser Antilles as far as Puerto Rico.
exfoliating in small strips, dark brown weathering
910 grey. Branches spreading, forming a wide crown in
Distribution
old trees. Foliage branches terete, finely grooved be-
tween leaf bases, terminating in small subglobose Caribbean: Trinidad.
buds with free, triangular, apiculate outer scales and TDWG codes: 81 TRT
imbricate inner scales. Leaves on shaded branch-
es and juvenile trees larger than on sun-exposed
Ecology
branches of mature trees, 6–10 cm long, 10–15 mm
wide, straight or sometimes slightly falcate. Leaves Podocarpus trinitensis occurs in montane, evergreen
on sun-exposed branchlets (2.5–)4–6(–7) cm long, tropical forest.
6–12 mm wide, spreading or directed forward on
ends of shoots, coriaceous, dark green above, pale
Conservation
green below, straight, elliptic to oblanceolate or ob-
long, gradually narrowing to a short petiolate base This species is indigenous to Trinidad, with an area
and more abruptly to an acute to weakly apiculate or of occupancy (AOO) of more than 20 km2. It does
rarely obtuse apex. Midrib on adaxial (upper) side not appear to be threatened or in decline, even
prominently raised but sometimes fading towards though it does not occur within a protected area.
apex, less than 0.5 mm wide and lying in a shallow IUCN: LC
depression, on abaxial side flat or forming a shallow
groove (in sicco). Stomata small, in numerous inter-
Uses
mittent, irregular lines on abaxial side. Pollen cones
axillary, solitary, sessile, subtended by thin, scarious No uses have recently been recorded of this species.
bud scales, cylindrical, maximum elongation not In the past, larger trees were probably used for con-
observed; microsporophylls triangular, minutely struction and general carpentry.
apiculate, bearing two relatively large, ovoid-oblong
pollen sacs. Seed cones axillary on short peduncles;
basal bracts (‘foliola’) deciduous; receptacles formed Podocarpus urbanii Pilg., in Engler, Pflanzenr.
of an axis with two unequal, fused bracts, swelling IV.5 [18]: 89. 1903. Type: Jamaica: Saint Thomas,
to a 7–8 × 6 mm large, succulent, red body. Seeds Blue Mountains, [“near the summit of the Blue
solitary, including the epimatium ovoid, 7–8 × 5–6 Mountains”], J. MacFadyen 27 (holotype K). Pl. 40
mm, with or without a distal, curved, thin crest.
Seed proper ovoid, smooth with a thin, light brown
Etymology
seed coat.
This species was named after the German bota-
nist Ignatz Urban (1848–1931), who published the
Taxonomic notes
first synthetic account of Caribbean or West Indies
There is some doubt about the taxonomic status of plants.
this species. The leaves are generally smaller than
those of P. coriaceus, but the dimensions appear to
Vernacular names
overlap, especially in younger trees or on branches
of shaded growth. The apex of leaves of P. trinitensis Yacca
 911

Pl ate 40. Podocarpus urbanii. 1, 2. Branches with leaves and seed cones. 3. Branchlet with leaves and
pollen cones. 4. Leaves. 5. Seed cone.
 with two fused, slightly diverging bracts, swelling
Description
to a 7–8 × 6 mm succulent, red ‘pseudo-fruit’. Seeds
Small trees to 16 m tall, often only 5–10 m. Bark solitary, including the epimatium obliquely ovoid,
smooth, dark reddish brown. Branches spreading, 6–7 × 4 mm with a distal conical crest, dark green
forming a wide crown in old trees. Foliage branches turning blackish. Seed proper not observed.
becoming densely leaved in mature trees with sun-
exposed crowns, terete, finely grooved between
Distribution
more remote leaf bases, terminating in small, glo-
bose buds with imbricate, keeled, obtuse or (outer Caribbean: Jamaica (Blue Mountains).
scales) minutely apiculate scales. Leaves on shaded TDWG codes: 81 JAM
912 branches and juvenile trees larger than on sun-ex-
posed branches of mature trees, 3–6 cm long, 5–9
Ecology
mm wide, straight or slightly falcate. Leaves on sun-
exposed branchlets (1–)2–3(–3.5) cm long, 3–5 mm Podocarpus urbanii occurs in montane to high mon-
wide, densely crowded and directed forward at an tane forests at altitudes between 1160 m and 2256 m
angle of 30–45° so as to obscure shoots, coriaceous, a.s.l. in the Blue Mountains of E Jamaica.
rigid, dark green above, straight, narrowly lanceo-
late; margins nowhere parallel, gradually converg-
Conservation
ing to a sessile base and an apiculate-pungent or
acute or rarely obtuse apex. Midrib on adaxial (up- In the Global Red List of Conifers (Farjon & Page,
per) side raised from leaf base to beyond the mid- 1999) this species was listed as Near Threatened
dle but fading towards apex, less than 0.5 mm wide, (LRnt following 1994 criteria) and this was still its
on abaxial side obscurely raised or flat or forming status in the IUCN Red List 2008. It has a limited
a shallow groove (in sicco). Stomata in two whitish distribution confined to a single mountain range,
bands of numerous irregular, wavy lines on abaxial where it is locally common. Reassessment seems
side. Pollen cones axillary, solitary, sessile, subtend- necessary for this species.
ed by thin, scarious bud scales, cylindrical, 12–15 IUCN: NT
mm long, ca. 4 mm wide; microsporophylls more or
less oval, obtuse with erose-scarious margins, bear-
Uses
ing two relatively large, ovoid-oblong pollen sacs.
Seed cones axillary on short peduncles; basal bracts No economic uses have been recorded of this spe-
(‘foliola’) deciduous; receptacles formed of an axis cies.
Prumnopitys Phil., Linnaea 30: 731. 1861. Type: Prumnopitys andina (Poepp. ex
Endl.) de Laub. (Podocarpus andinus Poepp. ex Endl.) [Prumnopitys elegans Phil.]
(Podocarpaceae).

Stachycarpus Tiegh., Bull. Soc. Bot. France 38: 163. 1a. Adult leaves spreading to all sides (but more
1891. Type: Stachycarpus andinus (Poepp. ex Endl.) or less pectinate on shaded shoots), slightly
Tiegh. [Prumnopitys andina (Poepp. ex Endl.) de curved backward or assurgent, slender, (1–)1.5–
Laub. (Podocarpus andinus Poepp. ex Endl.)] 2 mm wide P. taxifolia
1b. Adult leaves pectinately arranged, nearly
Greek: prumme, prumne = stern; pitys = pine or fir
straight or slightly curved sideways, sometimes 913
tree.
falcate, (1.8–)2–3.8 mm wide 2
2a. Small trees to 15 m tall, with a short bole or
Description
multi-stemmed. Adult leaves 1.8–2.5 mm wide.
Trees or multistemmed shrubs, evergreen, dioecious, Seeds including the fleshy epimatium 18–25 ×
with smooth or scaly bark. Resin canals in leaves 18–20 mm, bluish green ripening to yellow
(1) and resin in bark. Foliage branches alternate, P. andina
spreading in a more or less horizontal plane. Leaves 2b. Trees usually with a single bole and potentially
small, spirally inserted, in vegetative lateral shoots taller than 15 m. Adult leaves 2–3.8 mm wide.
usually pectinately arranged, flattened, more or less Seeds including the fleshy epimatium 10–20 ×
falcate-linear, epistomatic with stomata in two bands 7–15 mm, variably coloured (if larger than 20 ×
separated by a thin midrib. Pollen cones aggregate 15 mm, dark blue to blue-black) 3
on relatively long, leafless branchlets, cylindrical, 3a. Adaxial (upper) midrib (midvein) of leaves dis-
slender; pollen sacs relatively large, yellow, two on tinctly grooved. Pollen cones arranged in axil-
each microsporophyll, transversely dehiscent, con- lary spikes of 15–25 4
taining bisaccate pollen. Seed cones on small axillary 3b. Adaxial midrib of leaves indistinctly raised (not
branchlets with a few reduced leaves, consisting of a groove). Pollen cones solitary, nearly sessile 7
one to a few spirally arranged bracts developing a sin- 4a. Leaf apex obtuse, sometimes acute or acumi-
gle erect ovule (seed) in the axil of a small (terminal) nate. Seeds including the fleshy epimatium
bract, lacking any fleshy receptacle at maturity. Seeds crested, pruinose when ripe 5
ovoid, with a drupe-like, thick, fleshy and globose or 4b. Leaf apex acute or acuminate, sometimes mu­
ovoid-elliptic epimatium (outer layer) covering the cronate or obtuse. Seeds including the fleshy
seed entirely and becoming yellow, red or black. epimatium rounded or with an apical knob,
yellow or bluish, but not pruinose when ripe 6
9 species.
5a. Seeds (cones) on 0.5–1.5 cm long branchlets
with deciduous scale leaves. Microsporophylls
Distribution
of pollen cones with an acute apex, 0.5 mm
Australia: Queensland. SW Pacific: New Caledonia; wide P. exigua
New Zealand. America: Costa Rica (Central Ameri- 5b. Seeds (cones) on 1.5–2.5 cm long branchlets
ca); in South America along the Andes from W Ven- with persistent scale leaves at base and decidu-
ezuela to Bolivia; disjunct in S Chile. ous reduced leaves. Microsporophylls of pollen
cones with broadly triangular apex, 0.7 mm
wide P. harmsiana
Key to the species of Prumnopitys
6a. Seeds (cones) on 0.5–1.5 cm long branchlets
The vegetative characters are often very similar with a few reduced leaves, solitary. Seeds in-
among the species of Prumnopitys and the few dif- cluding the fleshy epimatium 10 × 7–8 mm,
ferent character states in the leaves that are easily ob- without an apical crest or knob P. standleyi
servable in the field are insufficient to differentiate 6b. Seeds (cones) on 2–3 cm long branchlets with
between them. Therefore pollen cones and ripe seeds (near) normal leaves, solitary or in pairs. Seeds
are necessary to arrive at a correct ­identification if including the fleshy epimatium 14–16 × 10–12
one has to make a choice from among all nine ­species. mm, with a distinct apical knob P. montana
 7a. Pollen cones very small, 3–4 × 1.3–1.6 mm. and young plants with slender leaves 8–15(–20) mm
Seeds including the fleshy epimatium 10–14 × long and 1.2–2 mm wide; apex acuminate. Leaves on
7–10 mm, with a ridge and apical knob mature trees alternate, twisted at a petiolate base,
P. ferruginoides mostly pectinately arranged in two rows and one
7b. Pollen cones larger, 7–20 × 2–3 mm. Seeds in- plane, linear, (8–)12–20(–25) mm long, 1.8–2.5 mm
cluding the fleshy epimatium 15–30 × 9–25 mm, wide, slightly curved sideways above base or more
only with an apical knob 8 or less falcate, with parallel sides, with a indistinct
8a. Adult leaves 2–3 mm wide, curved downward. proximal midrib adaxially (upperside) and a low
Pollen cones 8–20 mm long. Seeds including midrib abaxially (underside); margins entire; apex
the fleshy epimatium 15–18 × 9–13 mm, yellow acuminate or sometimes obtuse; leaf colour dull
914 becoming bright red P. ferruginea greyish green above, whitish green below. Stomata
8b. Adult leaves 2.5–3.5 mm wide, (nearly) straight. in two indistinct bands of numerous lines, more or
Pollen cones 7–9 mm long. Seeds including the less separated by midrib on abaxial (under) side of
fleshy epimatium 22–30 × 18–25 mm, dark blue leaves. Pollen cones on lateral and axillary spikes
to blue-black P. ladei towards ends of foliage branches; up to 25 spread-
ing cones per spike, 4–7 mm long and 2 mm wide,
each subtented by a leaf-like bract 3 mm long. Mi-
Prumnopitys andina (Poepp. ex Endl.) de Laub., crosporophylls sub-peltate with acute apex, 0.5 mm
Blumea 24 (1): 189. 1978. Podocarpus andinus wide, incurved at anthesis, light green, each with
Poepp. ex Endl., Syn. Conif.: 219. 1847. Type: two yellow pollen sacs. Seed cones 1–3 on 2–3 cm
[“Habitat in Chile australis alpinis frigidis, locis long branchlets with persistent scale leaves at base
umbrosis convallis Quillai-Leuvu versus Antuco. and deciduous, acute scale leaves, axillary to a scale
Pöppig msc.”] (HAL?, W destroyed). Fig. 296, 297 leaf (bract), consisting of an epimatium enclosing a
subglobose, slightly flattened, smooth seed, grow-
Podocarpus spicatus Poepp., in Poeppig, Nov. Gen. ing into a globose to broadly elliptical, bluish green
Sp. Pl. 3: 18. 1845, non R. Br. (1838); Prumnopitys spi­ ripening to yellow ‘pseudo-fruit’ 18–25 × 18–20 mm
cata (Poepp.) Molloy & Muñoz-Schick, New Zeal. diam. with an apical small knob.
J. Bot. 37: 190. 1999 (nom. illeg., Art. 53.3).
Prumnopitys elegans Phil., Linnaea 30: 732. 1860.
Distribution
Prumnopitys andina (Poepp. ex Endl.) de Laub. sub-
sp. blijdensteinii Silba, J. Int. Conifer Preserv. Soc. 9 S Argentina: Neuquén; S Chile: Biobio, La Arauca-
(1): 33. 2002. na, Maule.
TDWG codes: 85 AGS-NE CLC-BI CLC-LA CLC-MA
Etymology
Ecology
The species epithet refers to its occurrence in the
Andes. Prumnopitys andina occurs in valleys close to rivers,
in mixed forests with Austrocedrus chilensis, Cryp­
tocarya alba, Quillaja saponaria and other sclero-
Vernacular names
phyllous shrubs and trees. In the Andes and Coastal
Lleuque, Lleuqui, Uva de Cordillera (Chile) Cordillera it grows with Nothofagus dombeyi, N. ner­
vosa and N. obliqua in non-sclerophyllous montane
rain forests. Its altitudinal range is from 200 m to
Description
1400 m a.s.l.
Trees to 15 m tall; trunk to 1 m d.b.h., with a short bole
or multi-stemmed. First order branches spreading or
Conservation
ascending forming a broadly pyramidal or rounded
crown. Bark smooth on younger trees, becoming There are probably fewer than 12 subpopulations and
scaly and exfoliating in irregular small strips; bark only two of these have more than 1000 mature indi-
colour purplish grey, weathering grey. New foliage vidual trees, while many have fewer than 100. Several
glaucous green, turning greyish green. Seedlings subpopulations have been reduced recently, e.g. by
conversion of the natural forest to plantations of ex-
Description
otic trees for commercial forestry. Other threats are
clearing of woodland to make space for agriculture, Trees to 20 m tall; trunk to 1 m d.b.h., with a short,
flooding in relation to hydroelectric projects, and stout bole. First order branches spreading in large
increased wildfires. Regeneration is high in some trees forming a rounded crown. Bark smooth on
subpopulations but very low in others, possibly due younger trees, eventually exfoliating in irregular
to domestic animals eating the fleshy seeds. There are small papery strips; bark colour reddish brown,
three protected areas in Chile which contain this spe- weathering blackish brown. New foliage yellow-
cies: Reserva Nacional Nuble, Parque Nacional Con- ish green, turning dark green. Seedlings and young
guillio and Parque Nacional Tolhuaca. Germination plants with slender leaves 12–20 mm long and 1.8–
experiments conducted in Chile and the UK have re- 2.3 mm wide; apex acute. Leaves on mature trees 915
sulted in increased germination rates that could help alternate, twisted at a petiolate base, mostly pecti-
restocking lost or diminished natural populations. nately arranged in two rows and one plane, linear,
IUCN: VU [B2 a-b (ii-v)] 10–20 mm long, 2.2–3.1 mm wide, slightly curved
sideways above base, widest above the middle, with
a distinct grooved midrib adaxially (upperside)
Uses
and a midrib abaxially (underside); margins entire;
In the past the wood was much used to make furni- apex obtuse or sometimes acute; leaf colour lustrous
ture; at present trees are not felled at a commercial dark green above, dull light green below. Stomata in
scale but its wood can become available when for- two bands of numerous lines separated by midrib
est patches that contain it are cleared for other land on abaxial (under) side of leaves. Pollen cones on
uses. The succulent tissue surrounding the seeds is lateral and axillary spikes towards ends of foliage
sweet and these are harvested for consumption by branches; up to 20 spreading cones per spike, each
local people, as most female trees bear large quanti- cone axillary to a small leaf, 7–10 mm long and 2
ties of ‘fruit’ each year. There are also potential me- mm wide. Micro­sporophylls sub-peltate with acute
dicinal properties, e.g. chemicals that actively deter apex, 0.5 mm wide, incurved at anthesis, light green,
fungal infections detected in the bark of this tree. each with two yellow pollen sacs. Seed cones solitary
Prumnopitys andina is in cultivation in arboreta, bo- on 5–15 mm long branchlets with deciduous scale
tanic gardens and private tree collections in many leaves, axillary to a scale leaf (bract), consisting of
countries, mostly based on cuttings taken from ear- an epimatium enclosing a globose, slightly flattened
lier introductions. Growing anew from seed would seed, growing into a globose, pruinose ‘pseudo-fruit’
increase genetic diversity, thought to be crucial to 10–12 mm diam. with an apical crest.
counter potential pathogens.
Distribution
Prumnopitys exigua de Laub., Phytologia Mem. 7: Bolivia: Cochabamba, W Santa Cruz.
68. 1984. Type: Bolivia: Cochabamba, road from TDWG codes: 83 BOL
Sihuenca to Totora, M. Cardenas 4879 (holotype
US).
Ecology
This species occurs in upper montane evergreen
Etymology
rainforest. The altitudinal range (from information
The species epithet is from Latin: exiguus = short, on herbarium labels) is 1850 m to 3000 m a.s.l. At
insignificant; this species was at first believed to be a the upper limit this forest becomes stunted (low
stunted form, but larger trees were found later. cloud forest) and both vegetation formations are
rich in epiphytes. It appears to occupy rather similar
high altitude habitats as P. montana, but is disjunct
Vernacular names
from that species by ca. 1000 km.
pino colorado, pino negro (Spanish)
 dark brown, weathering blackish grey. Seedlings and
Conservation
very young plants with pectinately spreading leaves
Restricted to upper montane forest in two states in 7–15(–20) mm long and 1.4–2 mm wide. Leaves on
Bolivia. The extent of occurrence (EOO) is less than young trees alternate, twisted at a petiolate base,
20,000 km2 and there are 5–6 subpopulations in mostly pectinate, falcate, 15–30 mm long, 1.8–2.7
about 15 locations. A reduction in the extent of habi- mm wide, with a distinct, raised midrib on both
tat is caused by deforestation in the region, which sides; margins entire, tapering to an acute apex; leaf
affects this species. The area of occupancy (AOO) colour light green to brownish red. Leaves on adult
for this species has not been estimated but given the trees shorter, mostly pectinate and curved down-
15 localities known it probably falls below the 2000 ward, falcate-linear, 10–20 mm long and 2–3 mm
916 km2 threshold for Vulnerable. wide, with an indistinct midrib adaxially (above)
IUCN: VU [B1 a-b (iii)] and a distinct green midrib on the stomatal side;
apex obtuse or faintly apiculate; leaf colour lustrous
green above, whitish green below. Stomata in two
Uses
broad bands separated by a green midrib on abaxial
This species is probably logged with other podocarps (under) side of leaves. Pollen cones axillary, solitary,
if of sufficient size. The wood of this tree is valued nearly sessile, cylindrical, 8–20 mm long and 2.5–3
for construction and carpentry, but probably only of mm wide. Microsporophylls sub-peltate with apicu-
local significance. Many trees at higher altitudes are late apex, 1 mm wide, light green, each with two yel-
too stunted to be of any use for their timber, even low pollen sacs. Seed cones solitary on short axillary
though at least one old tree is known to have a trunk branchlets with deciduous scale leaves, subterminal
in excess of 1 m diam.; several herbarium collectors with 7–9 bracts but developing a single ovule (seed).
have taken their samples of it. It is not recorded to Epimatium enclosing a broadly ovoid, slightly flat-
be in cultivation. tened, smooth and ridged seed, growing into an el-
lipsoid, green or blue-green, then yellow and finally
bright red ‘pseudo-fruit’ 15–18 mm long and 9–13
Prumnopitys ferruginea (G. Benn. ex D. Don) mm wide, with a distinct apical knob.
de Laub., Blumea 24 (1): 190. 1978. Podocarpus
ferrugineus G. Benn. ex D. Don, in Lambert,
Distribution
Descr. Pinus, ed. 8°, 2: [189]. Type: “Habitat in
Nova Zelandia. Phillip, Bennett.” (LINN?, not New Zealand: North Island, South Island, Stewart
designated). Island.
TDWG codes: 51 NZN NZS
Etymology
Ecology
The species epithet means ‘with the colour of rusted
iron’ (Latin ferrum = iron) and refers to the leaves of Prumnopitys ferruginea is a constituent of podo-
young plants. carp forest occurring in lowland areas from near sea
level to as high as about 1000 m. Most of this for-
est has now gone and remnants are often disturbed
Vernacular names
or at any rate altered. Those forest reserves that still
Brown pine; miro (Maori) contain primary forest show a mixed podocarp-
broad-leaved forest with Dacrycarpus dacrydio­
ides, Dacrydium cupressinum, Podocarpus totara,
Description
Manoao colensoi, Prumnopitys taxifolia, and P. fer­
Trees to 25 m tall; trunk to 1 m d.b.h. First order ruginea as dominant conifers (all Podocarpaceae)
branches ascending and eventually spreading, in and a mixture of angiosperm trees and tree ferns in
old trees forming a domed or rounded crown. Bark a second layer. In Northland it also occurs with Aga­
more or less smooth on younger trees, covered in this australis (Araucariaceae). All of these are slow
tiny pustules, becoming scaly and exfoliating in growing (except for an initial stage of establishment)
thick flakes from the bole of large trees; bark colour and long-lived conifers with ‘K’ ecological ­strategies.
On the cooler west coast of the mountainous South
Description
Island, these conifers form a narrower belt below
Notho­fagus forest and are mixed with the south- Trees to 20 m tall; trunk to 0.5 m d.b.h., but of-
ern beeches in a transition zone. On Stuart ­Island ten remaining smaller, densely branched forming
mono­specific stands of P. ferruginea occur with a rounded crown. Bark more or less smooth on
Pseudopanax crassifolius in the understorey. younger trees, becoming scaly and breaking up into
small quadrangular flakes on the bole of large trees;
inner bark fibrous; bark colour yellowish or reddish
Conservation
brown, weathering grey. Leaves on young trees alter-
IUCN: LC nate, twisted at a short decurrent and more or less
petiolate base, mostly pectinate, (nearly) straight, 917
(10–)15–30 mm long, 3.5–4.5 mm wide, with a dis-
Uses
tinct, raised midrib on both sides; margins entire,
The wood of miro has similar properties as that of parallel to shortly below the obtuse or minutely
matai (P. taxifolia) and is used in construction for mucronate apex; leaf colour light green to brown-
floors and weatherboards. It sometimes is beauti- ish red. Leaves on adult trees much shorter, most-
fully figured, especially in radial cuts, which makes ly pectinate, linear or more or less oval in shortest
it desirable for furniture, e.g. table tops and cabinet leaves, (5–)8–12(–18) mm long and 2–3.8 mm wide,
making. Salmon (1996, repr. 2001) states that it is with a faint midrib on both sides; margins slightly
“exploited increasingly as a substitute for matai”, but incurved; apex obtuse or mucronate; leaf colour
that observation may be dated as the native timber lustrous green above, whitish green below. Stomata
producing conifer trees in New Zealand are now densely scattered on both sides of leaves, not sepa-
protected from logging. Perhaps this tree is used rated by midrib. Pollen cones axillary, solitary, on a 1
here and there under licence on a small scale, as it mm long peduncle, ovoid-oblong, 3–4 mm long and
is certainly more common and widespread than its 1.3–1.6 mm wide. Microsporophylls sub-peltate with
congener. It is rare in cultivation, restricted to some faintly apiculate apex, 0.5 mm wide, light green, each
botanical collections in warm temperate regions, with two yellow pollen sacs. Seed cones solitary on
e.g. California and Ireland. short axillary branchlets covered by persistent scale
leaves, terminal with 6–8 spreading, 3–4 mm long
bracts, but developing a single ovule (seed) subtend-
Prumnopitys ferruginoides (R. H. Compton) de ed by the distal, largest bract. Epimatium enclosing
Laub., Fl. Nouv. Calédonie Dépend. 4: 56. 1972. a broadly ovoid, slightly flattened seed, growing into
Podocarpus ferruginoides R. H. Compton, J. Linn. an ellipsoid, slightly flattened, green or blue-green
Soc., Bot. 45: 424. 1922. Type: New Caledonia: and finally dark blue ‘pseudo-fruit’ 10–14 mm long
Grande Terre, Province Sud, Mt. Nékando, and 7–10 mm wide, with a distinct polar ridge and
R. H. Compton 1073 (holotype BM). Fig. 298 apical knob.

Podocarpus distichus J. T. Buchholz, J. Arnold Arbor.

Distribution
32: 89. 1951.
Podocarpus distichus J. T. Buchholz var. maialis SW Pacific: New Caledonia.
J. T. Buchholz, J. Arnold Arbor. 32: 89. 1951. TDWG codes: 60 NWC

Etymology Ecology
The species epithet ferruginoides compares this spe- Prumnopitys ferruginoides is a tree of montane rain-
cies with P. ferruginea from New Zealand. forest distributed mainly on ultramafic soils derived
from serpentine; in the extreme north of the island
(Mont Panié) it occurs on micaschist. Its altitudinal
Vernacular names
range is approximately from 150 m to 1400 m a.s.l.
No vernacular names have been recorded for this This species is scattered in these forests as most-
species. ly solitary trees dispersed by birds, which eat the
 ‘pseudo-fruits’, and therefore this species can occur plane, linear, (8–)15–20 mm long, 2–3.5 mm wide,
randomly with many other trees, both conifers and slightly curved sideways above base, mostly paral-
angiosperms. Apparently this species is shade toler- lel-sided, with a distinct grooved midrib adaxially
ant, juveniles up to 5–6 m tall growing up under can- (upperside) and a thin midrib abaxially (underside);
opy have markedly larger leaves than mature trees of margins entire; apex mostly obtuse or slightly acu-
which the crowns have reached abundant sunlight. minate; leaf colour green above, glaucous below. Sto-
mata in numerous lines not separated by midrib on
abaxial (under) side of leaves. Pollen cones on lateral
Conservation
and axillary, 3–4 cm long spikes towards ends of fo-
IUCN: LC liage branches; 15–20 cones axillary to 2 mm long
918 bracts, 8–10 mm long and 2 mm wide. Microsporo­
phylls sub-peltate with broadly triangular apex, 0.7
Uses
mm wide, imbricate, light green, each with two yel-
There are no uses recorded of this species. If it is or low pollen sacs. Seed cones solitary on 1.5–2.5 cm
was logged it would have been treated as podocarp long axillary branchlets with persistent scale leaves
timber not distinct from other tree forming species, at base and deciduous reduced leaves, subtended by a
but its scarcity in the forest means it has no econom- longer scale leaf (bract); consisting of an epimatium
ic importance. As a rare tropical species it is not like- enclosing an ovoid, slightly flattened, rugose seed,
ly to be in cultivation outside a few botanic gardens. growing into a subglobose, pruinose ‘pseudo-fruit’
10 mm long and 8 mm wide with a small apical crest.

Prumnopitys harmsiana (Pilg.) de Laub., Blumea

Taxonomic notes
24 (1): 190. 1978. Podocarpus harmsianus Pilg.,
in Engler, Pflanzenreich IV.5 [18]: 68. 1903. Type: Pilger (op. cit.) mentioned collections by Karsten
Venezuela: Merida, Cordillera do Merida, Tovar, (unspecified) and A. Fendler (No. 1289) when de-
A. Fendler 1289 (lectotype K, designated here). scribing Podocarpus harmsianus as a new species
from Venezuela. When De Laubenfels (op. cit.)
Podocarpus utilior Pilg., Fedde’s Repert. Sp. Nov. transferred this species to the genus Prumno­pitys, he
Regni Veg. 1 (12): 189. 1905. did not designate its type; that opportunity is here
fulfilled with a duplicate of Fendler 1289 at K desig-
nated as the lectotype. It consists of a foliage branch
Etymology
of a probably juvenile (sterile?) treelet. Pilger saw
This species was named after the German botanist the spiked pollen cones, apparently on other mate-
H. A. T. Harms (1870–1942). rial which was likely destroyed at Berlin-Dahlem
(B) during World War II, and stated not to know the
seed cones. In describing Podocarpus utilior from
Vernacular names
Peru two years later, Pilger (op. cit.) had seed cones
uncumanu (Peru) but no pollen cones. His suspicion that the two
might be one species has been confirmed to be the
case and the two descriptions can be united.
Description
Trees to 35 m tall; trunk to 1.8 m d.b.h., with an erect
Distribution
bole. First order branches ascending or spreading in
large trees forming a rounded crown. Bark smooth The Andean regions of Bolivia, Colombia, Equador,
on younger trees, becoming scaly and exfoliating in Peru, and Venezuela.
irregular small strips; bark colour purplish brown, TDWG codes: 82 VEN 83 BOL CLM ECU PER
weathering grey. Seedlings and young plants with
slender, nearly straight to more or less falcate leaves
Ecology
20–30 mm long and 2–4 mm wide; apex acute. Leaves
on mature trees alternate, twisted at a ­petiolate base, Prumnopitys harmsiana is widespread but rela-
mostly pectinately arranged in two rows and one tively uncommon in the Andean lower montane to
­ ontane forest belt at altitudes between 1000 m and
m
Description
2200 m a.s.l.
Trees to 33 m tall; trunk to 1 m d.b.h., may be but-
tressed; upper part of tree densely branched form-
Conservation
ing a rounded crown. Bark more or less smooth on
This species has a wide distribution in a forest type younger trees, becoming scaly and breaking up into
that is under pressure throughout its range as a re- thin flakes on the bole of large trees; inner bark fi-
sult of deforestation. Most herbarium collections brous; bark colour reddish brown, weathering grey.
are from Bolivia and Peru. The extent of occurrence Leaves on young trees alternate, twisted at a short
(EOO) has not been calculated due to a lack of spec- decurrent and more or less petiolate base, mostly
imen locality data in the other countries, where it pectinate, (nearly) straight, 15–25 mm long, 3–4 mm 919
is believed to occur more frequently than the avail- wide, with a distinct, raised midrib on both sides;
able data suggest (data largely from the Tropicos margins entire, parallel to shortly below the obtuse
database, which does not include all relevant major and minutely hyaline apex; leaf colour green to
herbaria). The Conifer Specialist Group believes it brownish red. Leaves on adult trees shorter, mostly
is prudent to flag this species as NT as deforestation pectinate, linear or more or less oval in the short-
may well push it into a threatened category in fu- est leaves, (5–)8–12(–16) mm long and 2.5–3.5 mm
ture. This species has been recorded from the Machu wide, with a faint midrib on both sides; margins
­Pichu National Park in Peru. slightly incurved; apex obtuse or faintly apiculate;
IUCN: NT leaf colour green above, whitish green below. Sto-
mata densely scattered on both sides of leaves, not
separated by midrib. Pollen cones axillary, solitary,
Uses
sessile or on a short peduncle, oblong, 7–9 mm long
The timber of Prumnopitys harmsiana is valued for and 2–3 mm wide. Microsporophylls sub-peltate
the construction of houses and large trees are often with obtuse apex, 1 mm wide, light green, each with
selectively logged for that purpose. It is probably two yellow pollen sacs. Seed cones solitary on short
the tallest of the Andean species in the genus and, axillary branchlets with deciduous small leaves, ter-
depending on its forest setting, may grow a sizable minal with 1–2 bracts, the upper bract subtending a
straight bole. This species is not in cultivation out- single ovule (seed). Epimatium enclosing a broadly
side a few arboreta and botanic gardens in (sub) ovoid, slightly flattened brown seed, growing into an
tropical countries. ellipsoid to globose, green or blue-green and finally
dark blue to blue-black ‘pseudo-fruit’ 22–30 mm
long and 18–25 mm wide, with a distinct apical pro-
Prumnopitys ladei (F. M. Bailey) de Laub., Blumea trusion.
24 (1): 190. 1978. Podocarpus ladei F. M. Bailey,
Queensland Agric. J. 15 (8): 899, t. 22. 1905. Type:
Distribution
Australia: Queensland, [“Mount Spurgeon, Mitchell
River, Port Douglas, F. W. H. Lade, leaf specimens, Australia: NE Queensland, (Mt. Lewis, Mt. Spur-
December 1902; immature fruiting specimens, May geon and a few other localities).
1905.”] (syntypes, BRI). Fig. 299, 300 TDWG codes: 50 QLD-QU

Etymology Ecology
The species epithet commemorates F. W. H. Lade, This rare species is restricted to (remnants of)
who collected the type specimens in 1902 and 1905. rainforest habitat mainly on two mountains in NE
Queensland. These mountains are granitic outcrops
in the Atherton Tableland. The altitudinal range
Vernacular names
of the species is between 930 m and 1400 m a.s.l.
No common names have been recorded for this spe- The rainforest on these mountains is dominated by
cies. angio­sperms with scattered podocarps: Podocarpus
smithii, Prumnopitys ladei and Sundacarpus amarus.
 Prumnopitys ladei is the rarer of these species. The
Vernacular names
seeds are eaten by native rats.
diablo fuerte, pino criollo, pino de montana, romer-
illo hembra (Spanish)
Conservation
The surviving rainforests on both mountains are now
Description
partly within protected areas. Logging has almost
certainly reduced the area of occupancy (AOO) of Trees to 30 m tall; trunk to 1.5 m d.b.h., with an erect
Prumnopitys ladei on Mt. Lewis, presently the small- and straight bole, commonly buttressed at base. First
er of the two populations. In 1974 a herbarium col- order branches spreading or ascending in large trees
920 lection was made within the “North Mary Logging forming a broadly domed or rounded crown. Bark
Area” of State Forest Reserve 143 (B. P. M. Hyland smooth on younger trees, becoming scaly and exfo-
7882); this implies that logging of primary forest was liating in irregular small strips; bark colour purplish
then still undertaken and it would not have spared brown, weathering grey. New foliage yellowish or
this species. Several of these logging areas existed glaucous green, turning dark green. Seedlings and
there between 1960–1980 according to data on her- young plants with slender leaves 12–20 mm long and
barium labels. On the other hand, recent surveys 1.8–2.3 mm wide; apex acute. Leaves on mature trees
have extended the occurrence of P. ladei to a few lo- alternate, twisted at a petiolate base, mostly pecti-
calities in between both mountains. A reassessment nately arranged in two rows and one plane, linear,
of the conservation status of this species is required. (8–)12–20 mm long, 2–3.5 mm wide, slightly curved
IUCN: NT sideways above base, widest above the middle, with
a distinct grooved midrib adaxially (upperside) and
a midrib abaxially (underside); margins entire; apex
Uses
acuminate or sometimes obtuse; leaf colour lustrous
The wood of this species is used for construction and green above, glaucous below. Stomata in numerous
carpentry, but it is too rare to be of much economic lines not separated by midrib on abaxial (under)
importance. It is said to have potential as an orna- side of leaves. Pollen cones on leafless or nearly leaf-
mental tree but is rarely used in that capacity and less, lateral and axillary spikes towards ends of fo-
may not be present outside botanical collections. liage branches; up to 25 spreading cones per spike,
5–10 mm long and 2 mm wide. Microsporophylls
sub-peltate with acute apex, 0.5 mm wide, spreading
Prumnopitys montana (Humb. & Bonpl. ex Willd.) at anthesis, light green, each with two yellow pollen
de Laub., Blumea 24 (1): 189. 1978. Taxus montana sacs. Seed cones one or two on 2–3 cm long branch-
Humb. & Bonpl. ex Willd., Sp. Pl. 4 (2): 857. 1806; lets with leaves, axillary to a scale leaf (bract); con-
Podocarpus montanus (Humb. & Bonpl. ex Willd.) sisting of an epimatium enclosing an ovoid, slightly
Lodd. ex Britton, Bull. Torrey Bot. Club 16: 13. flattened, rugose seed, growing into an ovoid, olive-
1889. Type: [“Habitat in montibus Peruvianis.”] green ripening to yellow ‘pseudo-fruit’ 14–16 mm
(Humboldt & Bonpland in P-Bonpl.?, n.v.). long and 10–12 mm wide with an apical knob.

Podocarpus montanus (Willd.) Lodd. var. diversi­

Distribution
folius Dallim. & A. B. Jacks., Handb. Conif.: 51. 1923.
Podocarpus montanus (Willd.) Lodd. var. densifolius The Andean regions of Colombia, Equador, Peru
J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 70. and Venezuela.
1948. TDWG codes: 82 VEN 83 CLM ECU PER
Podocarpus montanus (Willd.) Lodd. var. meridensis
J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 71.
Ecology
1948.
Prumnopitys montana is a species of high montane
forest and timberline scrubland, ranging from 1500
Etymology
to 3600 m a.s.l. In the tropical high montane rain-
The species epithet refers to its habitat in mountains. forest (cloud forest) it is a constituent in a ­forest
type dominated by angiosperms such as Clusia rounded crown, foliage branches alternate, forming
(Guttiferae), Weinmannia (Cunoniaceae), Ocotea, flat sprays. Foliage twigs slender, glabrous, initially
and various other genera in Lauraceae. At the high- covered in narrow decurrent leaf bases, later with
est altitudes this forest is replaced by scrub, often thin, exfoliating bark. Leaves alternate, usually pec-
dominated by Ericaceae and Podocarpaceae, where tinately arranged in two rows and one plane, linear,
P. montana is an emergent but low tree or a dwarfed twisted and slightly curved above the decurrent base
tree. Both vegetation formations are rich in epi- and sometimes near apex, or more or less falcate,
phytes. In places with high water table near the tree (12)15–20(–25) mm long, 2–3.5 mm wide, narrowed
line Sphagnum bogs prevail, with the scrub vegeta- towards the petiolate base; apex acute to mucronate.
tion retreating to drier spots. Adaxial leaf surface (upperside) with a conspicuous
longitudinal groove through the middle, dark green; 921
abaxial surface (underside) with a narrow longitu-
Conservation
dinal midrib, with very small stomata in numer-
IUCN: LC ous lines covered with a greyish white wax. Pollen
cones numerous on slender, leafless, axillary spikes
towards ends of foliage sprays, close together, each
Uses
cone on a 2–3 mm long ‘stalk’ (bare base of the ra-
This podocarp is a very valuable timber tree; the chis), cylindrical, 10–15 mm long, 2–2.5 mm wide
wood is of medium density, straight grained, work- at anthesis. Microsporophylls numerous, helically
able and durable. It is used for construction, general arranged, sub-peltate, curved upwards with a scari-
carpentry, indoor floors, cabinet making, and tool ous denticulate, acute apex, each bearing near base
handles. It is also very good for wood turning with two relatively large pollen sacs. Seed cones terminal
light yellow sapwood and reddish heartwood. Where on small 0.5–1.5 cm long branchlets with a few re-
the forest has been cleared to make way for pasture, duced leaves; producing a single subapical, broadly
individual trees or small groups are often left stand- oval ‘pseudo-fruit’ ca. 10 mm long, 7–8 mm wide,
ing to provide shade for cattle. It is rare in cultiva- consisting of a fleshy epimatium enclosing a broadly
tion, mainly restricted to some botanic gardens. ovoid, slightly flattened seed, initially green, bluish
when mature.

Prumnopitys standleyi (J. T. Buchholz & N. E. Gray)

Distribution
de Laub., Blumea 24 (1): 190. 1978. Podocarpus
standleyi J. T. Buchholz & N. E. Gray, J. Arnold Costa Rica (Alajuela, Cartago, Heredia, San José).
Arbor. 29: 72. 1948. Type: Costa Rica: Alajuela, Poas TDWG codes: 80 COS
Mts., Volcan Poas, A. Tonduz 10333 (holotype US).
Ecology
Etymology
Prumnopitys standleyi is a climax forest species in
This species was named after the botanist Paul montane tropical rain forest. Its altitudinal range
C. Standley (1884–1963), who studied the flora of (GIS data) is 1349 m to 2603 m a.s.l.; based on her-
Costa Rica. barium specimen data it is 1800–2960 m, while
some reports mention altitudes above 3000 m. An-
nual rainfall is high, from 2000 mm to 4000 mm.
Vernacular names
The species is reported to be slow-growing.
Ciprecillo (Spanish).
Conservation
Description
This species is restricted to four to five locations.
Trees to 25(–30) m; trunk monopodial, to 1.5(–2) The extent of occurrence (EOO) based on specimen
m d.b.h. Bark smooth in young trees, becoming distribution data is calculated to be 611 km2 and the
scaly on large trunks, exfoliating in flakes, brown area of occupancy (AOO) 130 km2. Regeneration is
weathering grey. Branches spreading, forming a reported to be poor and trees are scarce in the forest,
 with 0.1–1 per ha according to one study. The tree ing to all sides but on some shaded branchlets more
is selectively logged, while only a few localities are pectinate, linear, (6–)10–20 mm long, (1–)1.5–2 mm
within protected areas. Some collections were made wide, slightly curved backward or assurgent, or
within the following protected areas: Volcan Poas nearly straight, with a distinct, slightly raised mid-
Reserve and Rio Macho Reserve. rib on both sides; margins entire; apex acuminate or
IUCN: EN [B1, B2 a-b (iii)] sometimes obtuse; leaf colour lustrous green above,
whitish green below. Stomata in two broad bands
separated by a green midrib on abaxial (under) side
Uses
of leaves. Pollen cones on leafless or nearly leafless,
The timber of Prumnopitys standleyi is valued for lateral and axillary spikes towards the ends of foliage
922 the construction of houses and large trees are of- branches; up to 25 spreading cones per spike, 5–12
ten selectively logged for that purpose. The wood is mm long and 2 mm wide. Microsporophylls sub-
suitable for construction, flooring, veneer, furniture peltate with rounded apex, 0.5 mm wide, light green,
and also used as fenceposts and for firewood. This each with two bright yellow pollen sacs, giving the
species is not in cultivation outside perhaps a few cones a yellow colour when mature. Seed cones sev-
botanic gardens. eral on stout branchlets with deciduous scale leaves,
axillary to a scale leaf (bract) and consisting of an
epimatium enclosing an ovoid, slightly flattened, ru-
Prumnopitys taxifolia (Banks & Sol. ex D. Don) gose seed, growing into a globose or nearly spheri-
de Laub., Blumea 24 (1): 190. 1978. Dacrydium cal, purplish black ‘pseudo-fruit’ 8–10 mm diam.
taxifolium Banks & Sol. ex D. Don, in Lambert, after most of the undeveloped, green seeds have
Descr. Pinus, ed. 1, 2: 25. 1824. Type: New Zealand: fallen.
[locality not stated], J. Banks & D. Solander s.n.
(holotype BM).
Taxonomic notes
The occurrence of pollen cones in spikes, i.e. situ-
Etymology
ated on what appear to be specialized axillary shoots
The species epithet refers to the yew-like leaves (yew without functional leaves and arranged in rows, is
= Taxus). rare in conifers. It occurs in Metasequoia, Taxodium
(both Cupressaceae) and in five species of Prumno­
pitys: P. andina, P. exigua, P. montana, P. standleyi,
Vernacular names
and P. taxifolia (Podocarpaceae). It is recorded from
Black pine; matai (Maori) the fossil record in Cordaitales, an extinct order of
gymnosperms that was perhaps ancestral to coni-
fers, and its limited occurrence in two not closely
Description
related extant conifer families indicates that this
Trees to 25 m tall; trunk to 1.3 m d.b.h., crown wide, character has evolved more than once. There may be
domed or rounded. First order branches ascend- some adaptive advantage, because these elongated
ing and eventually spreading in old trees open- shoots bearing pollen cones without leaves help to
ing up the wide spreading crown. Bark smooth on expose the former unimpeded to the air, so that the
younger trees, becoming scaly and exfoliating in slightest breeze could disperse the pollen.
irregular flakes leaving multi-coloured patterns on
the bole. Flakes on younger trees large, becoming
Distribution
much smaller on large boles of older trees, when
longitudinal grooves appear; bark colour pattern New Zealand: North Island, South Island.
varying from greenish grey or nearly white to red- TDWG codes: 51 NZN NZS
dish or purplish brown. New foliage bright green,
turning dark green. Seedlings and young plants with
Ecology
long, whip-like shoots and brownish leaves 5–10
mm long and 1–2 mm wide. Leaves on mature trees Prumnopitys taxifolia is a constituent species of low-
alternate, twisted at a petiolate base, mostly spread- land to montane (20–800 m or even 1000 m a.s.l.)
podocarp (conifer) forest, dominated by Podo­carpus
Conservation
totara, Dacrycarpus dacrydioides, or Dacrydium
cupressinum, and with Prumnopitys spp., Manoao IUCN: LC
colensoi, Halocarpus kirkii, and Phyllocladus tri­
chomanoides as frequent associated conifers. In low-
Uses
land areas few undisturbed forest remains and here
numerous angiosperms are often mixed in, while at Matai is a slow growing tree and produces dense,
higher altitude especially in South Island’s Westland hard and heavy brown wood with exceptional
District the genus Nothofagus is often codominant. strength and durability. It was extensively logged
As a slow growing conifer, reaching a maximum and used for railway sleepers, bridges, house build-
age in excess of 1000 years (Enright & Hill, 1995) ing, especially floors and weatherboards, and to a 923
Prumno­pitys taxifolia is a species that typically re- lesser extent for furniture. The logging of this and
generates only after episodal distrurbances such other native conifers is now outlawed and conse-
as fires caused by volcanic eruptions. It then com- quently the use of its wood has greatly diminished.
petes with other invading podocarps as well as an- It is rare in cultivation and only suitable for regions
giosperms for light and space, resulting in the long with mild winters that rarely experience light frosts
term survival of only a few individuals of this spe- and have plentiful and evenly distributed rainfall.
cies in the forest.
 Pseudolarix Gordon, Pinetum: 292. 1858 (nom. cons.). Type: Pseudolarix amabilis
(J. Nelson) Rehd. [Larix amabilis J. Nelson] (Pinaceae).

Laricopsis Kent, in Veitch, Man. Conif., ed. Kent: Pseudolarix pourtetii Ferré, Trav. Lab. Forest. Tou-
403. 1900 (nom. illeg.). Type: Laricopsis kaempferi louse T. 1 (4, 4): 1, f. 1–13. 1944.
(Lindl.) Kent [Abies kaempferi Lindl.]. Chrysolarix
H. E. Moore, Baileya 13: 133. 1965. Type: Chrysolarix
Etymology
amabilis (J. Nelson) H. E. Moore [Larix amabilis
J. Nelson]. The species epithet (Latin amabilis = ‘lovely’ or ‘lov-
924 able’) refers to its golden yellow autumn foliage.
Greek: pseudo = false; i.e. resembling but not equal-
ling the genus Larix.
Vernacular names
Description Chinese golden larch; jin qian song (Chinese)
See the species description.
Description
Taxonomic notes
Monoecious, deciduous trees to 30–40 m tall, d.b.h.
The genus Pseudolarix is known from the fossil record to 1.5 m; trunk monopodial, straight or curved;
(Early Cretaceous to Pleistocene) in numerous lo- crown irregular, broadly domed and open in old
calities in North America and Eurasia. It occurred in trees. Bark rough and scaly, breaking into thick,
Europe at least until the end of the Pliocene and in square scales, brown-grey. Branches spreading hori-
the Early Tertiary extended far north into the Arctic. zontally or slightly ascending, drooping at ends.
At least two species were present during the Tertiary. Branchlets slender, flexible, in second year thicker
It is a classic example of a relict conifer which once and firm, pale yellowish, darkening to pinkish or
was very common and widespread, but has been re- purplish brown, later grey, ridged and grooved, gla-
duced to a few localities in China. This conifer was brous, young shoots slightly pruinose. Short shoots
‘traditionally’ classified with Cedrus and Larix based cylindrical-oblong, marked with annual rings of
on vegetative characters but more detailed phylo- perular scale bases, length increasing with age, 0.5–
genetic analyses using both morphology and DNA 3.5 cm. Vegetative buds of long shoots ovoid, pale
have refuted this. Its affinities are with Keeteleria, brown, (axillary) lateral buds of short shoots globu-
Nothotsuga and Tsuga in an ‘abietoid’ clade (Farjon, lar, yellowish green, 2–3 mm, not resinous. Leaves
1990; Wang et al., 2000) separate from a ‘pinoid’ clade spirally arranged, remote on long shoots, dense on
which includes among others Cedrus and Larix. The short shoots, in false terminal whorls of 10–30, as-
short shoots and deciduous leaves are parallelisms surgent, (2–)3–6(–7.5) cm long, (1.5–)2–3(–4) mm
which evolved at least twice within the Pinaceae. wide, linear, flaccid, flattened, shallowly grooved
above, keeled below, obtuse or acutish at apex;
Distribution hypo­stomatic, stomata in 2 broad bands separated
by a midrib; leaf colour light green, pale green be-
As for the species.
low, turning bright yellow, finally orange-brown
in ­autumn. Pollen cones terminal on short shoots,
Pseudolarix amabilis (J. Nelson) Rehd., J. Arnold Ar- clustered with 10–25 in umbels, ovoid at first, cylin-
bor. 1: 53. 1919. Larix amabilis J. Nelson, Pinaceae: 84. drical and pendulous when ripe, becoming yellow.
1866; Chrysolarix amabilis (J. Nelson) H. E. Moore, Seed cones terminal on short shoots, solitary, erect,
Baileya 13: 133. 1965. Type: United Kingdom: England, with short-leaved, 0.5–1.5 cm long peduncles, ovoid-­
cultivated, G. Gordon s.n. (holo­type K). Fig. 301, 302 globose, open, resembling artichokes, (4–)5–7(–8)
cm long, 4–5.5 cm wide with opened scales, col-
Abies kaempferi Lindl., Penny Cyclop. 1: 34. 1833, our light green or purplish green, ripening to light
(quoad descr., non basion.); Pseudolarix kaempferi yellowish brown, disintegrating; cone rachis weak,
(Lindl.) Gordon, Pinetum: 292. 1858. narrowly conical. Seed scales narrowly deltoid-
triangular, auriculate-pedicellate at base, slightly tends to angiosperm (broad-leaved) dominance.
recurved or straight at apex, 2.8–3.5 × 1.3–1.8 cm Undisturbed primary forest that contains P. amabilis
at mid-cone, coriaceous, striated or rugose at ma- is now extremely rare and, if it still exists, should be
turity, sometimes with orange-yellow resin, nearly subjected to detailed ecological study.
glabrous; margins entire, erose in old scales. Bracts
lanceolate, 6–8 mm long, entirely included. Seeds
Conservation
ovoid-cuneate, partly enveloped by wing membrane,
5–7 mm long, pale brown or rose-brown; seed wings The natural distribution of this species is based upon
obliquely pointed, extending beyond the seed scales, the populations in and around Zhejiang and two
23–30 × 8–13 mm, yellow or pale brown. localities in southern Anhui and northern Jiangxi
and (tentatively) one in Hunan; all other herbarium 925
based localities are believed to be planted or intro-
Distribution
duced and here and there perhaps naturalized; they
China: lower Chang Jiang (Yangtse) valley (Anhui, are excluded from the assessment. This species is
Hunan, Jiangxi, Jiangsu, Zhejiang). The true geo- very rare in the wild and occurs in a few remnants
graphic range of this species is difficult to ascer- of primary forest on isolated mountains. Most loca-
tain. In Atlas of the Gymnosperms of China (Ying tions are not within protected areas and loss of habi-
et al., 2004), a map with dots (based on herbarium tat is still continuing in this densely populated part
specimen data from Chinese herbaria) is produced of China. A detailed survey of existing truly natural
which gives a much wider range than here present- populations occurring in primary mixed forest to
ed. A map given in China Plant Red Data Book 1 determine more exactly the number of remaining
(Fu & Jin, 1992) is similarly optimistic. A check by mature trees, as well as the number of populations
Dr Xiang Qiao-ping in the Beijing Herbarium (PE) and their localities, is a first and necessary step to
revealed that most of these specimens were collected conservation action. This should next lead to the es-
from planted trees or trees in villages and towns (e.g. tablishment of protected areas including the major
near temples) that did not occur in the wild. This sites and populations.
problem is acknowledged in Flora of China 4 (1999). IUCN: EN [B1a, b (iii)]
Wang (1961) has given an indication where we may
expect this attractive tree to occur in its natural hab-
Uses
itat: his examples are all in northern Zhejiang.
TDWG codes: 36 CHS-AH CHS-HN CHS-JS CHS-JX This species has been planted in arboreta and parks
CHS-ZJ in many countries, providing ample material for
study. Herbarium specimens from (wild) locations
in China are very rare, therefore the majority of ma-
Ecology
terial studied has come from European arboreta.
Pseudolarix amabilis is a component of the mixed Due to its rarity and slow growth the value as a tim-
mesophytic, partly evergreen forest, very rich in tree ber tree of this conifer is limited to local use, pri-
species (Wang, 1961). Its altitudinal range is from 180 marily for boat building, wooden foot bridges, and
m to 1000 m a.s.l., so it is a component of lowland furniture. The Chinese golden larch has been in cul-
forest in a warm temperate, humid climate which tivation in Europe since the early 1860’s when it was
can experience occasional cold winter frost at the first described as a species of larch in England, based
higher altitude. This species grows on a variety of on small plants grown from seed and from seedlings
soils derived from acidic rock; it does not occur on transported by Robert Fortune from China to Eng-
limestone. Several localities where P. amabilis still land in a Wardian case. It is a desirable and success-
occurs in the wild are likely to have been altered by ful tree in cultivation, which has been responsible
man and will support secondary forest, indicated by for its wide ‘distribution’ in China probably centuries
the presence of Cunninghamia lanceolata and ­Pinus before it came to Europe. It is now present in most
spp.; some of these trees may have been planted in collections of planted conifers in the temperate cli-
reforestation projects nearby. On the other hand, mate regions of the world, especially in the northern
P. amabilis is a light demanding tree and perhaps not hemisphere, but it remains relatively uncommon in
a constituent of a late phase in the succession, which the trade, perhaps because it is ­difficult to propagate
 and slow growing. It is mostly used as a specimen
tree in large gardens and parks in W Europe and
the eastern USA. A few dwarfed forms have been
selected and are grown as cultivars in Chinese and
Japanese style gardens.

926
Pseudotaxus W. C. Cheng, Notes Forest. Inst. Nat. Centr. Univ. Nanking, Dendrol.
Ser., 1: 1. 1947. Type: Pseudotaxus chienii (W. C. Cheng) W. C. Cheng [Taxus chienii
W. C. Cheng] (Taxaceae).

Nothotaxus Florin, Acta Horti Berg. 14 (9): 394. 1948 bushy crown. Foliage branchlets subwhorled or
(nom. illeg.). Type: Nothotaxus chienii (W. C. Cheng) subopposite, slender, terete, smooth, green turn-
Florin [Taxus chienii W. C. Cheng]. ing dark green in the second year, terminating
in small, ovoid-conical buds with persistent bud
Greek: pseudo- = false, i.e. resembling but not equal- scales. Leaves helically arranged, distichous, spread-
ling; Taxus is the classical Latin name for yews. ing at 40–90° to shoot axis, well spaced but some 927
overlapping at various angles, 1.2–2.8 cm long, lin-
ear, straight or slightly falcate in some leaves only,
Description
(1.5–)2–4(–4.5) mm wide, short petiolate or ses-
See the species description. sile, obtusely rounded at base; margins parallel and
revolute, abruptly converging and terminating in a
mucronate apex; mucro hooked, pale whitish green;
Distribution
leaf texture slightly coriaceous, smooth or slightly
As for the species. rugose; leaf colour green above, abruptly turning
dark green in second year, green with two white
bands below. Midrib raised on both sides, ca. 0.4
Pseudotaxus chienii (W. C. Cheng) W. C. Cheng, mm wide, continuous. Stomata in two bands 0.5–1
Notes Forest. Inst. Nat. Centr. Univ. Nanking, mm wide on abaxial (lower) side, slightly wider than
Dendrol. Ser., 1: 1. 1947. Taxus chienii W. C. Cheng, midrib and green margins, forming 10–20 intermit-
Contr. Biol. Lab. Sci. Soc. China, Sect. Bot. 9: 240. tent white lines in each band. Pollen cones usually
1934; Nothotaxus chienii (W. C. Cheng) Florin, Acta concentrated towards ends of lateral branchlets,
Horti Berg. 14 (9): 394. 1948. Type: China: Zhejiang, axillary, solitary, sessile, globose, 2–3.5 mm long,
Longquan Xian, Maoshan, S. Chen 1384 (lectotype 3 mm wide, with 4 pairs of decussate basal bracts;
PE). Fig. 303 microsporophylls 6–12, decussate, peltate, with 4–6
radially placed pollen sacs. Seed-bearing structures
Pseudotaxus liana Silba, Phytologia 81 (4): 327. axillary, solitary, sessile, with 7 pairs of decussate
1996 [“liiana”]; Pseudotaxus chienii (W. C. Cheng) basal bracts. Seed arils cupular, only partly enclos-
W. C. Cheng subsp. liana (Silba) Silba, J. Int. Conifer ing the seed, succulent and white when ripe, 5–7
Preserv. Soc. 14 (1): 19. 2007. mm long. Seeds ovoid, 5–8 mm long, 4–5 mm wide,
slightly flattened distally, with two obscure ridges
below mucronate apex.
Etymology
The species epithet commemorates S. S. Chien, a
Taxonomic notes
Chinese botanist who was Head of the Botanical
Division of the Biological Laboratory of the Science This is a monotypic genus endemic to SE China.
Society of China. Pseudotaxus liana was described by Silba (op. cit.)
from a specimen collected in Guangxi and said to
have slightly broader leaves which are “leathery”,
Vernacular names
but the leaf measurements at least appear to overlap
Whiteberry yew; bai dou shan (Chinese) considerably with P. chienii as described in Flora of
China 4 (1999). In a recent morphometric study of
herbarium specimens in Chinese institutions (very
Description
little material of this taxon is available in Western
Dioecious, evergreen shrubs to 4 m tall. Bark herbaria), Wang & Yang (2007), although find-
grey-brown, exfoliating in strips. Branches usu- ing statistically significant differences in mean leaf
ally whorled, spreading or ascending, forming a width between specimens from Zhejiang (P. ­chienii)
 and from outside that province, also found that is infrequent, with seed maturation from late Sep-
these different measurements among the specimens tember to October and seeds remain dormant for a
are continuous. They therefore reduced P. liana to year or more.
synonymy of P. chienii, a conclusion which is here
accepted.
Conservation
This species, although distributed widely, is very rare
Distribution
and Endangered due to deforestation. Fragmenta-
China: N Fujian, N Guangdong (Nan Ling Shan, Ru- tion of populations into individual shrubs affects
yian), Guangxi, N and SW Hunan, SW Jiangxi, SW pollination success, which is already naturally dif-
928 Zhejiang. ficult for an understorey species in evergreen forests.
TDWG codes: 36 CHS-FJ CHS-GD CHS-GX CHS-HN According to Fu & Jin (1992) this taxon has suffered
CHS-JX CHS-ZJ rapid decline in recent times due to destruction of
habitat. Some localities are within nature or forest
reserves, e.g. in the Fenyang Shan and Jiulong Shan
Ecology
(Zhejiang), the Jinggang Shan (Jiangxi), the Nan
Pseudotaxus chienii occurs scattered in evergreen and Ling Shan (Guangdong), and the Zhangjiajie Shan
deciduous broad-leaved forests of middle altitude (Hunan). Urgent further protection is needed, both
mountains in warm temperate regions. The climate in situ and ex situ; some efforts are being made in
on these mountains is temperate, cool and humid, China to cultivate the species.
with annual precipitation around 1800–2000 mm IUCN: EN (A1c)
and frequent cloud cover or fog. It grows in shallow
yellow earth (pH 4.2–4.5), or in crevices on acidic
Uses
rock, often on precipices above streams. In Zhejiang
it is associated with Schima superba, Acer elegatulum The wood is used as a carving material and for mak-
and Rhododendron simianum. In Guangxi it is found ing utensils. The species has been planted in the
with Schima argentea and Castanopsis eyrei, always botanic garden of Nanjing, Fujian, and is in cultiva-
in the shade of these trees. In N Guangdong and tion as an ornamental on a limited scale in Zhejiang.
SW Hunan I have seen Pseudotaxus chienii growing Outside China this interesting species is not in cul-
over rock precipices above streams or roads. With tivation, although W. C. Cheng introduced it in the
exposure to sun its growth becomes retarded and 1930s to the Arnold Arboretum in Massachussetts,
the habit changes to a low shrub. Seed production where plants grew for some time before they died.
Pseudotsuga Carrière, Traité Gén. Conif., ed. 2, 1: 256. 1867. Type: Pseudotsuga
menziesii (Mirb.) Franco [Abies menziesii Mirb.] (Pinaceae).

Abietia A. H. Kent, in Veitch, Man. Conif., ed. Kent:

Key to the species and some varieties of
474. 1900. Type: Abietia douglasii (Sabine ex D. Don)
Pseudotsuga
A. H. Kent [Pinus douglasii Sabine ex D. Don].
1a. Seed scales not wider than long, thin; bracts
Greek: pseudo = false, i.e. resembling but not equal- much longer than the seed scales, usually not
ling the genus Tsuga. reflexed. Leaves 1–1.5 mm wide, parted above
the shoot but not pectinate. Bark on the lower 929
trunk of large trees very thick and deeply fis-
Description
sured P. menziesii
Monoecious, sometimes very tall (–100 m), ever- 1b. Seed scales wider than long, relatively thick;
green trees, monopodial with a columnar trunk and bracts as long as or slightly longer than the seed
a conical or spreading crown and pseudo-whorled, scales, often reflexed. Leaves 1.5–2 mm wide,
plagiotropic branching (Massart’s model). Resin mostly pectinate. Bark on the lower trunk rela-
canals in wood, bark, leaves and seed cones. Bark tively thin 2
thick, with deep, longitudinal fissures in large trees. 2a. Seed cones ovoid-cylindrical, very large (9–18
(Sub-)terminal buds ovoid conical, not resinous, cm long); bracts slightly longer than the seed
with triangular scales persisting 1–2 years. Leaves scales, usually not reflexed. Leaf apex acute
spirally arranged, spreading more or less radially or P. macrocarpa
pectinate, placed on small depressions, linear, flat- 2b. Seed cones ovoid, smaller than 8 cm; bracts
tened, with two bands of stomata only on the abaxial equally long as the seed scales, reflexed. Leaf
side. Pollen cones axillary, solitary, catkin-like, 1–2 apex usually emarginate 3
cm long, subtended by conspicuous perular scales; 3a. Seed cones broad, ovoid, usually 4.5–7.5 × 4–5.5
microsporophylls spirally arranged, peltate, with cm. Leaves variable in length, up to 5 cm long
two pollen sacs containing globular pollen with (leaves 0.7–2 cm long and 2–3 mm wide = P. sin­
3 converging ridges on a smooth surface. Seed cones ensis var. brevifolia) P. sinensis var. sinensis
axillary, solitary, more or less erect at pollination but 3b. Seed cones smaller (not longer than 5 cm).
soon pendulous on curved peduncles, deciduous Leaves up to 3 cm long 4
some time after seed dispersal or semi-persistent. 4a. Seed cones ovoid-conical; seed scales variable,
Bract scales exserted, as long as or longer than the slightly wider than long, with rounded upper
seed scales, trilobate, with the cusp longer than the margin P. japonica
lateral lobes, straight or reflexed. Seed scales with a 4b. Seed cones ovoid-oblong; seed scales reniform
rounded upper margin and a broad base, opening to broad flabellate, much wider than long, with
more or less wide. Seeds held in a shallow membra- truncate upper margin
nous cup covering one side of the seed and which is P. sinensis var. gaussenii
continued in an adnate, short, obliquely ovate wing.
Seedlings with (5–)7–10(–14) cotyledons.
Pseudotsuga japonica (Shiras.) Beissn., Mitt.
4 species. Deutsch. Dendrol. Ges. 1896 (5): 62. 1896. Tsuga
japonica Shiras., Bot. Mag. (Tokyo) 9: 84, t. 3. 1895.
Type: Illustration in Bot. Mag. (Tokyo) 9, t. 3 opp.
Distribution
p. 41. 1895. (lectotype designated here). Fig. 304, 305
North America: from SE Alaska (coast) and British
Columbia to Pueblo, Mexico in coastal mountains
Etymology
and the North American Cordillera, southward in-
creasingly scattered. Asia: in the eastern Himalaya, The species epithet refers to Japan, its country of ori-
Central and SE China, N Vietnam, Taiwan. gin.
 TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Vernacular names
Japanese Douglas-fir; toga-suwara, goyo-toga (Japa-
Ecology
nese)
Pseudotsuga japonica occurs at elevations between
500 m and 650 m a.s.l. on Shikoku; Wilson (1916)
Description
reports it from about 1000 m on the same island. It
Trees to 25–30 m tall, d.b.h. to 1–1.5 m; trunk usually is rare and scattered in sheltered valleys and steep
monopodial, more or less straight. Bark breaking ravines, growing on old volcanic rock. The climate
into vertical, flaking plates, in old trees longitudi- is warm temperate, moist, with annual precipitation
930 nally fissured, grey. Branches spreading out in all di- between 1000 and 2000 mm. It is locally a (major)
rections, crooked; branches of second order slender, constituent of mixed conifer-broad-leaved forests,
spreading horizontally or ascending; crown broad, with Tsuga sieboldii usually as the dominant spe-
open, usually domed or flat topped. Branchlets cies. Other conifers are Abies firma, Chamaecyparis
slender, firm, yellowish grey, becoming light grey- obtusa, Torreya nucifera and Cryptomeria japonica,
ish brown or grey, glabrous, with small, angular leaf broad-leaved trees are e.g. Quercus salicina, Q. ses­
scars. Vegetative buds ovoid to fusiform-conical, silifolia, Cleyera japonica, and Illicium religiosum
acute, 4–6 × 2–3 mm, not resinous; bud scales trian- (small tree), common shrubs are Eurya japonica,
gular, older scales with erose margins, shining red- Pieris japonica, Rhododendron serpyllifolium, and
dish brown, persisting 1–2 years. Leaves spreading Thea japonica. As in China, in Japan the genus, rep-
irregularly on both sides of the shoot, or pectinate, resented by a single species, is a minor constituent
especially on shaded vegetative shoots, (1.5–)2–3 cm of mixed forests.
long, 1.5–2 mm wide, slightly twisted or curved at
base, linear, straight, longitudinally grooved above,
Conservation
flattened, (slightly) emarginate at apex; stomata
in two bands separated by a midrib below (abax- The total population of this species in Japan is esti-
ial side); leaf colour (light) green above, two whit- mated to be ca. 2000 mature trees and is considered
ish stomatal bands below. Pollen cones 1–1.5 cm to be continuously declining.
long, yellowish brown. Seed cones on 1–2 cm long, IUCN: VU (C2)
curved peduncles, ovoid or ovoid-conical, taper-
ing towards apex when closed, 3.5–5.5(–6) cm long,
Uses
2–3.5(–4) cm wide with opened scales, pale purplish
or purplish brown, ripening to dark, dull brown or Japanese Douglas fir is of minor importance as a tim-
blackish brown; old cones deciduous after shedding ber tree due to its rarity and modest size, although
seeds, but often persisting 2–4 years thereafter. Seed veteran trees can attain substantial trunk diameters.
scales flabellate to semi-orbicular or sub-reniform, The wood is used for construction of traditional
convex, 1.5–2.2 × 2.5–3 cm at mid-cone; upper mar- buildings and gateways and for furniture. In Japan,
gin entire, incurved; base shortly narrowed. Bracts it may be seen planted in some parks and traditional
oblong-spathulate, with 3-lobed apex, central cusp gardens, outside its country of origin it is a rare tree
longer and narrower than lateral lobes, 1.5–2 cm only present in some arboreta. This species was col-
long, widest (–8 mm) near apex, exserted, recurved lected by Ernest H. Wilson in 1914 and introduced
or reflexed. Seeds ovoid to more or less triangular, via the Arnold Arboretum of Harvard University to
4 × 7 mm, pale brown, with dark spots; wings ovate, the USA. In Europe it seems to have arrived a few
4–10 mm long, light or dark brown. years earlier, perhaps a little after 1900 in Germany.
It grows slowly in cultivation and seems to remain a
small tree with a rather limited life span.
Distribution
Japan: W Honshu (Chugoku District), Kyushu,
Shikoku.
Pseudotsuga macrocarpa (Vasey) Mayr, Wald. r­ eflexed. Seeds ovoid-conical, 10–12 × 6 mm, brown;
Nord­amer.: 278, t. 6, 8, 9. 1889. Abies macrocarpa wings obovate, 10–14 × 6–8 mm, light brown.
Vasey, Gard. Monthly & Hort. 18: 21. 1876; Pseudo­
tsuga menziesii (Mirb.) Franco subsp. macrocarpa
Distribution
(Vasey) E. Murray, Kalmia 12: 24. 1982. Type not
designated. USA: S California (San Rafael Mts., San Gabriel
Mts., San Bernardino Mts., San Jacinto Mts., and
Santa Ana Mts.).
Etymology
TDWG codes: 76 CAL
The species epithet means ‘with large fruit’ and re-
fers to the seed cones. 931
Ecology
Vernacular names Pseudotsuga macrocarpa grows on the seaward
slopes of coastal mountains at elevations between
Bigcone Douglas-fir
275 m and 2450 m a.s.l., on mountain soils of vari-
ous origin, usually rocky and well drained, dry in
Description
summer. The climate is warm temperate, with cool,
Trees to 20–25 m tall, d.b.h. to 1–1.3 m; trunk mo- moist winters and warm, dry summers (annual pre-
nopodial, straight. Bark on trunk dark, blackish cipitation 500 mm to 1500 mm), snow occurs only
grey, scaly, longitudinally fissured, exposing reddish at the higher elevations. Above 800 m (Transition
brown inner bark. Branches spreading horizontally, Zone) this species is mainly mixed with Pinus jef­
the lowest may be bent down; crown broad pyrami- freyi, P. ponderosa, P. coulteri, and Abies concolor at
dal, very open in old trees. Branchlets firm when the highest elevations. Common evergreen shrubs
short, slender, flexible and pendulous when longer, in these open pine forests are Ceanothus cordulatus
reddish brown or pale brown, in second year grey- and Arctostaphylos patula. At the lower elevations
brown, faintly ridged, minutely pubescent when it occurs in the drier Canyon Live Oak forest type,
young, soon glabrous; leaf scars small, circular. Vege- with sclerophyllous oaks, e.g. Quercus kelloggii,
tative buds ovoid-conical to fusiform-conical, acute, Q. chrysolepis and Q. agrifolia.
7–8 × 4–5 mm, not resinous; bud scales triangular,
with serrulate-erose margins, reddish brown, per-
Conservation
sisting 1–2 years. Leaves directed forward and side-
ways, several raised above shoot, on shaded shoots The range of this species is relatively small and con-
more pectinate, 2.5–3.5(–4) cm long, ca. 2 mm wide, fined to the Transverse and Peninsular Ranges of
more or less petiolate and strongly twisted at base, the State of California, USA. The long, hot summers
linear, straight, longitudinally grooved above, flat- make it susceptible to forest fires, which are by far
tened; apex acute; stomata in two grey-white bands the main threat to this no longer exploited species.
separated by a midrib below; leaf colour lustrous It has limited resistance to fire, but as fire frequency
dark green or glaucous green above. Pollen cones has increased due to human causes, protection from
2 cm long, pale yellow, with red brown perular scales forest fires has become a major concern. As yet there
at base. Seed cones becoming pendulous on 1–2 cm seems to be no marked decrease of total known area
long peduncles and deciduous, ovoid-cylindrical, of occupancy (AOO) since surveys on the geo-
more or less pointed at apex, 9–13(–18) cm long, 4–6 graphic relationships between this conifer and wild-
cm wide with opened scales, greenish yellow, with fires were first reported in 1980, probably thanks to
light green bracts, maturing to light brown, ripening fire fighting and preventive measures.
to dull brown, with light brown bracts. Seed scales IUCN: NT
broadly cuneate-flabellate, thick, transversely con-
vex, 2–2.5 × 3–3.5 cm at mid-cone, puberulent when
Uses
young, soon glabrous, often resinous; upper margin
rounded, entire; base cuneate-pedicellate. Bracts The wood of this species is close grained, hard and
ligulate, with trilobate apex and a central cusp long- heavy, but not durable. It would only be suitable for
er than lateral lobes, 3–3.5 cm long, exserted, not coarse lumber, but is not exploited due to scarcity of
 the resource and other, more ecological values. It is cm long, 2–4 cm wide with opened scales, green,
perfectly suitable for amenity planting and as a spec- with yellowish green bracts, ripening to brown or
imen tree in gardens and parks in milder climates, dull grey brown, with light brown bracts. Seed scales
but rarely seen outside specialist collections in arbo- obovate-cuneate, convex, 2–2.5 × 2–2.5 cm at mid-
reta and botanic gardens. It grows to a well-shaped, cone; base more or less cuneate. Bracts ligulate, tri-
medium sized tree with attractive, large, pendulous lobate at apex, with cusp longer than lateral lobes,
seed cones and is worth being introduced and plant- 2.5–3.5 cm long, exserted, not reflexed, often curved
ed more often; obviously in regions with a suitable towards cone apex. Seeds ovoid-cuneate, 6–8 × 4–5
climate, such as southern Europe, or perhaps New mm, light brown, often with dark spots; wings ovate
Zealand and parts of Australia. oblong, 9–14 × 5–8 mm, yellowish brown.
932
Distribution
Pseudotsuga menziesii (Mirb.) Franco, Conif.
Duarum Nom.: 4. 1950 [& Bol. Soc. Brot., ser. 2, 24: W North America: from British Columbia to Cen-
74. 1950]. tral Mexico (Puebla).
TDWG codes: 71 ABT BRC 73 COL IDA MNT ORE
WAS WYO 76 ARI CAL NEV UTA 77 NWM TEX 79
Etymology
MXC-PU MXE-CO MXE-CU MXE-DU MXE-HI MXE-
The species was named after Archibald Menzies, SL MXE-ZA MXN-SO
who sailed with George Vancouver along the west
coast of North America in 1794.
Ecology
Pseudotsuga menziesii occurs in a huge area from
Vernacular names
north to south (55° N to 19° N) in W North America,
Douglas-fir, Oregon pine (wood) consequently it occupies a variety of climatic zones,
landscapes and habitats. Along the coast in British
Columbia and the Pacific Northwest this species at-
Description
tains great size and is a codominant or dominant
Trees to 90–100(–110?) m tall, d.b.h. to 4–5 m; trunk tree in the temperate rainforests from near sea lev-
monopodial, straight, columnar. Bark on large trunks el to 1000 m a.s.l. depending on latitude. It profits
rough and scaly, with deep, longitudinal fissures, from the high rainfall, yet occupies better drained
dark grey brown or blackish grey. Branches spread- sites commonly on slopes or high, no longer flood-
ing horizontally, but lower ones curved downward; ed river terraces, where it can compete successfully
crown of young trees broad conical, sometimes with with other conifers, mainly Abies, Picea, Tsuga, and
drooping leader, in old trees more columnar or ir- Thuja, especially after fire. Giant trees attain 100
regular. Branchlets slender, flexible, at first yellowish m, and there is evidence of taller trees that were
green later grey, minutely pubescent in the first year, logged in the days that Americans of European de-
then glabrous, leaf scars small, circular. Vegetative scent could see only the timber in them. Somewhat
buds ovoid-conical, acute, ca. 10 × 5 mm, not resin- further inland the species grows also in valley bot-
ous; bud scales triangular, lustrous reddish brown, toms near streams, still attaining great sizes and liv-
persisting 1–2 years. Leaves spreading more or less ing to 800–1000 years maximum. These coniferous
radially, on shaded shoots more pectinate, (1.5–)2.5– forests are of similar composition as those on the
3.5(–4) cm long, 1.2–1.5(–1.7) mm wide, more or less coast. In the Rocky Mountains occurs var. glauca, a
petiolate and slightly twisted at base, linear, straight, smaller, but still large tree; here it occupies a mixed
flattened, obtuse or acute at apex; stomata in two conifer belt between open Pinus contorta and/or
whitish bands separated by a midrib on abaxial (= P. ponderosa woodland and a subalpine conifer for-
under) side, lustrous dark green above or more or est dominated by Abies lasiocarpa, Picea engelman­
less glaucous. Pollen cones 1.5–2 cm long, yellow, nii or, further south, Pinus albicaulis and P. flexilis.
with red brown perular scales at base. Seed cones In the southern Rocky Mountains and into Mexico,
pendulous on 0.5–1 cm long peduncles, deciduous, Pseudotsuga menziesii var. glauca becomes more and
ovoid-conical, obtuse or pointed at apex, 4–9(–10) more scattered and restricted to sites with perma-
nent moisture, e.g. under N-facing canyon walls and TDWG codes: 71 BRC 73 ORE WAS 76 CAL
at the highest forested altitudes, up to 2900–3350
m a.s.l. In canyons on the Colorado Plateau it can
Conservation
form small groves rising above Gambel oaks (Quer­
cus gambellii), or it occurs as a constituent of mixed IUCN: LC
conifer forests with Pinus spp. and sometimes Abies
or Picea as well as Aspen (Populus tremuloides) on Pseudotsuga menziesii (Mirb.) Franco var. glauca
high plateaus and on N-facing slopes and in moist (Beissn.) Franco, Conif. Duarum Nom.: 6. 1950 [&
ravines at high altitude. Winter snowfall, unlike on Bol. Soc. Brot., ser. 2, 24: 76. 1950]. Tsuga douglasii
the Pacific coast, constitutes a high proportion of (Sabine ex D. Don) Lindl. var. glauca Beissn., in
annual precipitation in these habitats. Jäger & Beissner, Ziergeh. Gärt. Park., ed. 2: 446. 933
1884; Pseudotsuga menziesii (Mirb.) Franco subsp.
glauca (Beissn.) E. Murray, Kalmia 12: 24. 1982.
Uses
Type not designated.
Douglas fir is one of the world’s most important tim-
ber trees. The huge size, especially of the coastal va- Pseudotsuga flahaultii Flous, Bull. Soc. Hist. Nat.
riety, as well as the excellent wood properties make Toulouse 66: 332. 1934; Pseudotsuga menziesii
it a choice tree providing knot-free sawn timber of (Mirb.) Franco var. flahaultii (Flous) Silba, Phytolo-
great length. It is used for plywood and construc- gia 68: 70. 1990.
tion, both exterior and interior, and its reasonable Pseudotsuga menziesii (Mirb.) Franco var. oaxacana
good durability makes it useful for telephone wire Debreczy & Rácz, Phytologia 78 (3): 21. 1995.
poles and railway sleepers. However, the more con-
tinental variety P. menziesii var. glauca grows much
Description
slower and to a more moderate size and yields dens-
er, heavier wood, excellent for cooperage for vats This variety remains a smaller tree than P. men­
and tanks for breweries and distilleries. Douglas fir ziesii var. menziesii and has usually shorter, wider
has been introduced to many countries in plantation (1.5–2.5(–3.5) cm × 1.4–1.7 mm) and more glaucous
forestry as well as an ornamental tree and a good green leaves. The female cones are usually smaller,
number of cultivars are known and used in horticul- but variable.
ture. Its first introduction was in England in 1827 by
David Douglas; its best growth is attained in western
Distribution
Scotland, where one tree has made nearly 65 m in
about 100 years. In the NW USA and W Canada it W North America: Rocky Mountains from British
is also grown as a Christmas tree, needing regular Columbia to Central Mexico (Puebla).
shearing to obtain a pyramidal shape. TDWG codes: 71 ABT BRC 73 COL IDA MNT WAS
WYO 76 ARI NEV UTA 77 NWM TEX 79 MXC-PU
2 varieties are recognized:
MXE-CO MXE-CU MXE-DU MXE-HI MXE-SL MXE-
ZA MXN-SO
Pseudotsuga menziesii (Mirb.) Franco var. menzi­
esii. Abies menziesii Mirb., Mém. Mus. Hist. Nat. 13:
63, 70. 1825. Type not designated. Fig. 306, 307, 308 Conservation
IUCN: LC
Description
Potentially very tall trees to 100 m and more. Leaves
(2–)2.5–3.5(–4) × 1.2–1.5 mm, dark green. Seed cones Pseudotsuga sinensis Dode, Bull. Soc. Dendrol.
4–9(–10) cm long. France 23–24: 58. 1912.

Distribution Etymology
W North America: from British Columbia to Cali- The species epithet refers to China.
fornia.
 Vernacular names Xizang [Tibet], Yunnan, Zhejiang; Taiwan; N Viet-
nam.
Chinese Douglas-fir; huang shan (Chinese)
TDWG codes: 36 CHC-GZ CHC-HU CHC-SC CHC-
YN CHN-SA CHS-AH CHS-FJ CHS-GX CHS-HN CHS-
Description
JX CHS-ZJ CHT 38 TAI
Trees to 50 m tall, d.b.h. to 2 m; trunk more or less
straight, columnar, or forked. Bark on trunk rough
Ecology
and very scaly, longitudinally fissured below, grey.
Branches spreading wide, nearly erect near the top, Pseudotsuga sinensis is a species occurring in low to
forming a domed or flat topped crown. Branchlets medium high mountains at various elevations. In SE
934 slender, reddish brown in the first year, soon becom- China it occurs between 600 m and 1200 m, in Taiwan
ing grey, variably but usually minutely pubescent between 1000 and 2700 m, in Sichuan and Yunnan
in the grooves, soon glabrous, with small, slightly it may be found above 3000 m, the highest record is
elevated, circular or angular leaf scars. Vegeta- 3300 m a.s.l. The soils in the SE are red and yellow
tive buds ovoid or ovoid conical, 4–8 × 3–4.5 mm, earth, in the western part of the range described as
not or only slightly resinous; bud scales triangular, mountain red earth (Wang, 1961). It requires a moist,
acute, lustrous red-brown, deciduous in the second temperate or warm temperate climate with annual
year. Leaves more or less remote, pectinate, (0.7–) precipitation between 1000 and 2000 mm. It is a con-
2.5–4(–5) cm long, highly variable, (1–)1.5–2.1(–3) stituent of the mixed mesophytic forest formation in
mm wide, slightly twisted or curved at base, lin- SE China, mainly with broad-leaved trees, in Sichuan
ear, usually straight, longitudinally grooved above, also of the evergreen oak and deciduous hardwood
flattened, emarginate or sometimes obtuse at apex; forest, where besides Pseudotsuga other conifers oc-
hypostomatic, stomata in two greenish white bands cur (e.g. Tsuga chinensis, T. dumosa, Picea brachytyla
separated by a midrib on abaxial (= under) side; leaf var. complanata). Unlike in North America, Pseudo­
colour dark green above. Pollen cones pendulous tsuga in Asia does not form extensive stands or oc-
at maturity, 1–1.5 cm long, yellowish. Seed cones cur in pure or nearly pure conifer forests. Its habit
on 1–2.5 cm long peduncles, persisting some years as a mature forest tree reflects this: trees are not co-
but finally deciduous, ovoid to ovoid-oblong, (3.5–) lumnar or pyramidal but develop spreading crowns
4–6.5(–8) cm long, (2–)3.5–5(–5.5) cm wide with much like the surrounding broad-leaved trees.
opened scales, purplish, maturing to purplish brown
or brown. Seed scales rhombic-orbicular, suborbic-
Uses
ular, or semi-orbicular to reniform, convex, usually
apically slightly thickened and lignified, 2.5–3 × 3–3.5 Chinese Douglas fir is a timber tree used for con-
cm at mid-cone, slightly puberulent at first, but soon struction, bridge building, furniture and wood fiber.
glabrous; upper margin entire; base short pedicel- Very large trees are rare as they have been extensively
late. Bracts ligulate-linear, with trilobate apex, cusp logged in the past; hence the economic value of this
longer than lateral lobes, 3.5–4 cm long, exserted, re- species has diminished as it appears not very suit-
curved or reflexed. Seeds cuneate-ovoid, 8–12 × 5–8 able for plantation forestry. Although introduced in
mm, light brown with dark spots; seed wings ovate, Europe under its various synonyms by E. H. Wilson,
8–15 mm long, brown with dark spots. George Forrest, Camillo Schneider, and others in
the early 20th century, the species remains very rare
in cultivation as an amenity tree and is virtually re-
Taxonomic notes
stricted to a few arboreta and botanic gardens.
In Flora of China 4: 38 (1999) P. brevifolia and P. for­
3 varieties are recognized:
restii are treated as distinct species and P. wilsoniana
is recognized as a variety of P. sinensis.
Pseudotsuga sinensis Dode var. sinensis. Type:
China: Yunnan, Dali Baizu Zizhizhou, Dengchuan,
Distribution E. E. Maire s.n., 1911 (holotype P). Fig. 309, 310
China: S Anhui, Fujian, SW Guangxi, Guizhou, W Pseudotsuga wilsoniana Hayata, Icon. Pl. Formos. 5:
Hubei, N Hunan, N Jiangxi, S Shaanxi, Sichuan, SE 204, t. 15. 1915; Pseudotsuga sinensis Dode var. wil­
soniana (Hayata) L. K. Fu & Nan Li, Novon 7 (3):
Taxonomic notes
263. 1997.
Pseudotsuga forrestii Craib, Notes Roy. Bot. Gard. Pseudotsuga sinensis var. brevifolia has been reported
Edinburgh 11: 189. 1919; Pseudotsuga sinensis Dode from the limestone region in northern Vietnam, but
var. forrestii (Craib) Silba, Phytologia 68: 72. 1990. more recent assessments of the Vietnamese trees (e.g.
Pseudotsuga xichangensis C. T. Kuan & L. J. Zhou, Fl. Nguyen Tien Hiep et al., 2004) have concluded that
Sichuan. 2: 54. 1983. the common species on the ridge tops and summits of
Pseudotsuga shaanxiensis S. Z. Qu & K. Y. Wang, the limestone mountains is P. sinensis with normally
Acta Bot. Bor.-Occid. Sin. 8 (2): 129. 1988. long leaves. I confirmed this on a visit to Bat Dai Son
in 2002 at least for that area. The distribution of P. sin­
ensis var. brevifolia in Atlas of the Gymnosperms of 935
Description
China (Ying et al., 2004) on pp. 100–101 borders the
Leaves 1.3–4 cm long, 1.5–2 cm wide. Buds ovoid- distribution of P. sinensis in Vietnam given on a map
conical, more or less acute, 4–7 × 3–4 mm. Seed in Nguyen Tien Hiep et al. (2004). Is var. brevifolia
cones 4.5–8 cm long, 3–5 cm wide when opened. restricted to China? It seems unlikely, but it may also
Seed scales suborbicular to rhombic-orbicular. be incorrect determination of specimens on either
side of the border that has lead to this ‘parapatric’
distribution pattern. A critical evaluation of the ma-
Distribution
terial (and this taxon) seems needed.
China: S Anhui, Chongqing, Fujian, Guizhou,
W Hubei, N Hunan, N Jiangxi, S Shaanxi, Sichuan, Distribution
Yunnan, Zhejiang; Taiwan.
China: SW Guangxi, Guizhou (Anlong, Lipuo); N
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
Vietnam (?).
SC CHC-YN CHN-SA CHS-AH CHS-FJ CHS-HN CHS-
TDWG codes: 36 CHC-GZ CHS-GX
JX CHS-ZJ 38 TAI
Ecology
Conservation
Scattered on S-facing slopes and mountain tops, on
A continuing decline, especially of large mature trees calcareous and rocky soils at altitudes between 1000
due to logging, has led to a situation in which large and 1300 m a.s.l.
trees have become scarce and most are restricted to
inaccessible mountain ridges and summits. Even
Conservation
under the broader taxonomic circumscription here
employed, it has become a rare, though still wide- This variety occurs in a limited area within the ex-
spread tree. Only small subsets of the total popula- tensive eroded limestone massifs of SW China; its
tion are within protected areas. true extent and rarity remains unknown (see under
IUCN: VU [B1, 2b (iii, v)] Taxonomic notes). Trees are often inaccessible on
mountain summits but those on easier sites have of-
Pseudotsuga sinensis Dode var. brevifolia (W. C. ten already been logged.
Cheng & L. K. Fu) Farjon & Silba, Phytologia 68: 71. IUCN: VU (B1+2a-b)
1990. Pseudotsuga brevifolia W. C. Cheng & L. K. Fu,
Acta Phytotax. Sin. 13 (4): 83. 1975. Type: China: Pseudotsuga sinensis Dode var. gaussenii (Flous)
Guangxi, Nanning, near Longzhou, Chinese collec­ Silba, Phytologia 68: 71. 1990. Pseudotsuga gausseni
tor No. 40103 (holotype PE). Flous, Bull. Soc. Hist. Nat. Toulouse 69: 417. 1936.
Type: China: Zhejiang, between Ping-yung and
Taiyuan, R. C. Ching 2144 (holotype NY).
Description
Leaves 0.7–1.5(–2) cm long, 2–3(–3.2) mm wide.
Description
Buds ovoid, obtuse, 4–5 × 3 mm. Seed cones 3.7–6.5
cm long, 3–4 cm wide when opened. Seed scales Leaves 1.6–3 cm long, 1.7–2 mm wide. Buds ovoid-
suborbicular to rhombic-orbicular. conical, more or less acute, 4–5 × 3 mm. Seed cones
 3.5–5.5 cm long, 2–3.5 cm wide when opened; seed
Conservation
scales broad flabellate to reniform.
A continuing decline especially of large mature
trees due to logging has led to a situation in which
Distribution
large trees have become scarce and most are re-
China: SE Anhui, Fujian (?), Jingxi (Dexing), Si- stricted to inaccessible mountain ridges and sum-
chuan, N & NW Zhejiang. mits. IUCN: VU (B1+2c)
TDWG codes: 36 CHC-SC CHS-AH CHS-JX CHS-ZJ

936
Retrophyllum C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: 379. 1989.
Type: Retrophyllum vitiense (Seem.) C. N. Page [Podocarpus vitiensis Seem.]
(Podocarpaceae).

Decussocarpus de Laub., J. Arnold Arbor. 50: 340. 2b. Adult leaves (6–)10–25(–40) mm long, 2.5–5(–
1969 (nom. illeg., Art. 52.1). Type: Decussocarpus 7) mm wide; midrib on the adaxial (upper) side
nagi (Thunb.) de Laub. [Myrica nagi Thunb.]. of leaves inconspicuous or wide and low 3
3a. Midrib on both sides of leaves wide and low.
Latin: retro- = backwards, reversed; Greek: phyllos Microsporophylls of pollen cones triangular,
= leaf; referring to the peculiar phyllotaxis (see de- acute R. comptonii 937
scription). 3b. Midrib inconspicuous on the adaxial (upper)
side of leaves, narrow and conspicuous on the
abaxial side. Microsporophylls of pollen cones
Description
triangular, acuminate 4
Dioecious, evergreen, dwarfed to large trees. Resin 4a. Adult leaves (6–)10–18(–25) mm long. Pollen
canals in leaves and seed cones. Leaves spirally in- cones 5–7 × 2–2.5 mm. Seeds including the epi-
serted on shoots, lanceolate to narrowly ovate, ob- matium 18–25(–30) × 13–18(–20) mm
liquely directed into subopposite and decussate R. rospigliosii
apparent pairs by at ca. 90° twisted petioles in op- 4b. Adult leaves (10–)15–25(–40) mm long. Pollen
posite directions, forming regular pectinate rows on cones 10–25 × 2–2.5 mm. Seeds including the
lateral shoots. Stomata on both faces of the leaves. epimatium 14–20 × 10–13 mm R. vitiense
Pollen cones axillary or sometimes terminally and
single or in groups on short, naked peduncles; pol-
len bisaccate. Seed cones axillary, solitary or some- Retrophyllum comptonii (J. T. Buchholz)
times in pairs, at maturity consisting of a few small, C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45:
fused basal bracts and a single large seed covered 380. 1989. Podocarpus comptonii J. T. Buchholz,
by a fleshy, drupe-like, elliptic to ovoid-pyriform Bull. Mus. Hist. Nat. (Paris), sér. 2, 21: 284. 1949;
epimatium colouring deep red, violet or purplish Decussocarpus comptonii (J. T. Buchholz) de Laub.,
brown. J. Arnold Arbor. 50: 344. 1969; Nageia comptonii
(J. T. Buchholz) de Laub., Blumea 32 (1): 211. 1987.
5 species. Type: New Caledonia: Grande Terre, Province Sud,
Mt. Mou, J. T. Buchholz 1684 (holotype ILL). Fig.
311, 312
Distribution
Malesia: Maluku [Moluccas], New Guinea, New
Etymology
Britain, New Ireland; SW Pacific: New Caledonia,
Vanuatu, Fiji. South America: W Venezuela, Colom- This species was named after the botanist and plant
bia, Equador, Peru, W Brazil. collector R. H. Compton (1886–1979).

Key to the species of Retrophyllum Vernacular names

1a. Shrubs or small, dwarfed trees; foliage branches No vernacular names are recorded for this species.
ascending or spreading. Leaf apex always ob-
tuse R. minus
Description
1b. Potentially tall trees; foliage branches spreading
or more or less pendulous. Leaf apex acute, api- Trees to 30 m tall, to 80 cm d.b.h., bole straight and
culate or some obtuse 2 erect, clear of branches for 10 m or more in large trees
2a. Adult leaves (5–)8–11 mm long, 2–3.5 mm wide; growing in forest. Bark smooth, eventually becom-
midrib on both sides of leaves narrow and con- ing rough and fissured, tan to dark brown, weather-
spicuous R. piresii ing grey, exfoliating in longitudinal flakes. Branches
 ascending or spreading, forming a pyramidal and forested slopes and ridges of the mountains through-
eventually rounded crown. Foliage branches alter- out the main island (Grande Terre). It occurs on
nate or subopposite, spreading but becoming pen- serpentine and other ultramafic rocks as well as on
dulous in shaded branches, with remotely dispersed, acidic micaschist (e.g. Mont Panié). It is shade toler-
decurrent, spreading, small and thick scale leaves on ant and grows up well amongst other trees; it is not
leading shoots and at base of lateral shoots and more an emergent, but moves into the prevalent canopy,
numerous and larger, photosynthetically dominant which can be up to 30 m tall in sheltered spots, but is
leaves only on lateral branchlets. Larger leaves on often much lower, especially on mountain ridges and
lateral branchlets spirally inserted, appearing oppo- summits. Common associated conifers are Araucar­
site, pectinate on young plants and shaded shoots, ia spp. and Agathis spp. (Araucariaceae), Acmopyle
938 more or less spreading in different directions on sun pancheri, Falcatifolium taxoides, and to a lesser ex-
exposed branchlets, with a decurrent base and a 90° tent Podocarpus spp. (Podocarpaceae); there are also
twist at the point where the leaf blade abruptly wid- numerous angiosperms in these forests. Its altitudi-
ens and is free from the shoot, twisting in opposite nal range is between 600 m and 1450 m a.s.l. Dis-
directions, orientating the blade alternatively with persal by birds accounts for a scattered occurrence
adaxial side uppermost and with adaxial side down- of mature trees from where seedlings can spread in
wards. Leaf shape ovate-lanceolate or ovate-elliptic, the surrounding vegetation closer to the parent tree.
longest in juvenile plants, much shorter in top of old
trees, also shorter at the base of lateral branchlets, Conservation
the free part (6–)13–24(–32) × 2.5–4(–6) mm, mostly
IUCN: LC
widest below the middle, especially juvenile forms
gradually tapering to an acute, or in short leaves
Uses
obtuse apex; midrib wide but low on both sides.
Stomata on both sides (leaves amphistomatic), nu- This species grows into a large tree in favourable
merous and scattered, not separated by the midribs. habitat and has been logged with other trees espe-
Pollen cones grouped at end of foliage branchlets, on cially in lowland forest. The wood has good timber
lateral non-foliar branchlets, or solitary in the axil qualities and is used for light construction, flooring
of a leaf, globose when immature, becoming short and furniture. Trees on upper slopes and mountain
cylindrical, 5–10(–12) mm long and 2.5–3 mm diam. ridges are generally too small or short stemmed for
at anthesis; microsporophylls spirally arranged, tri- timber harvesting. This species is not in cultivation
angular, with an acute apex, each bearing two pollen except as some recently collected young specimens
sacs. Seed cones on 10–15 mm long lateral or ter- in a few botanic gardens.
minal branchlets with 2–3 pairs of decussate scale
leaves, with a single (rarely two) fertilized ovule
growing into an inverted seed covered by an epima- Retrophyllum minus (Carrière) C. N. Page, Notes
tium. Seeds including the epimatium obovoid, with Roy. Bot. Gard. Edinburgh 45: 380. 1989 [“minor”].
a curved beak at inverted apex and a short crest at Nageia minor Carrière, Traité Gén. Conif., ed. 2,
distal end usually disappearing at maturity, 16–20 × 2: 641. 1867; Podocarpus minus (Carrière) Parl., in
13–15 mm, green or glaucous becoming fleshy and Candolle, Prodr. 16 (2): 509. 1868; Decussocarpus
deep red when ripe; seed proper subglobose, 13–17 × minus (Carrière) de Laub., J. Arnold Arbor. 50: 346.
10–12 mm, with shallow scallops and ridges. 1969. Type: New Caledonia: Grande Terre, Province
Sud, Plaine des Lacs [“Baie du Sud”], E. Vieillard
1275 (holotype P). Fig. 313, 314
Distribution
SW Pacific: New Caledonia. Podocarpus palustris J. T. Buchholz, Bull. Mus. Hist.
TDWG codes: 60 NWC Nat. (Paris), sér. 2, 21: 284. 1949.

Ecology Etymology
Retrophyllum comptonii is a tree forming species of The species epithet minus denotes the small stature
montane rainforests, occurring commonly on steep of this tree, it being the smallest in the genus.
 Vernacular names Taxonomic notes
bois bouchon (French) Establishing the type specimen of R. minus poses
some difficulty in that the geographical information
for Vieillard 1275, the type collection, differs on vari-
Description
ous sheets distributed to different herbaria. Carrière
Small trees or shrubs to 3 m tall, with a short, thick (op. cit.) mentions “Lac Arnaud” in the protologue,
trunk quickly tapering upwards to short, stout this appears on the Paris (P) specimen, but other
branches, or sometimes multi-stemmed. Bark very specimens (BM, K) have “Baie du Sud” as the local-
thick on trunk, fibrous, soft, exfoliating in thin ity. Neither of these toponyms are in current use and
strips, brown (often discoloured by iron oxide were at any rate written on the labels later by some- 939
from the river in which it stands). Branches few, one who was not the collector. There are also male
spreading, forming an open, irregular crown. Foli- and female shoots mounted on different sheets with
age branches alternate or subopposite, ascending or No. 1275, which seems to be a number Vieillard add-
spreading, often grouped together towards ends of ed to his specimens after they were collected (some-
main branches, stout, subtended by a few spread- times by someone else; one sheet at K has “Pancher
ing scale leaves. Leaves on lateral branchlets spirally No. 1275” as the collection of a female shoot with
inserted, appearing opposite, distichous on juvenile separate seed). Since the species is dioecious, these
plants, more or less spreading in different direc- different sexes were collected from different plants.
tions on mature plants, with a decurrent base and a Carrière only described vegetative characters and it
90° twist at the point where the leaf blade abruptly is assumed that he only had the male specimen at
widens and is free from the shoot, twisting in oppo- P at his disposal, but did not notice the small pol-
site directions, orientating blade alternatively with len cones. Isotypes therefore are limited to male (or
adaxial side uppermost and with adaxial side down- sterile) specimens and female specimens are not
wards. Leaf shape ovate-lanceolate or ovate-elliptic, type material. The locality information on the labels
longest in juvenile plants, shorter in mature plants, seems to be of less significance than the appearance
in the latter those set near base of lateral branchlets that the sterile/male shoots all seem to have been
very small, free part (3–)10–20(–30) × 2–4(–6) mm, gathered from the same plant.
mostly widest in the middle, juvenile forms more or
less linear-lanceolate, all leaves with an obtuse apex;
Distribution
midrib wide but low on both sides. Stomata on both
sides (leaves amphistomatic), numerous and scat- SW Pacific: New Caledonia (Plaine des Lacs).
tered, not separated by the midribs. Pollen cones 2–5 TDWG codes: 60 NWC
clustered at end of foliage branchlets, or solitary in
axil of a leaf below these, globose when immature,
Ecology
becoming short cylindrical, 4–8 mm long and 2.5–3
mm diam. at anthesis; microsporophylls spirally This is the only true rheophyte among gymno-
arranged, triangular, with an apiculate apex, each sperms. It grows along the banks of small rivers and
bearing two pollen sacs. Seed cones terminal on shores of small lakes in shallow water over skeletal
­foliage branches ending in short branchlets with 2–3 soils derived from serpentine, ironstone and laterite
pairs of decussate scale leaves, with a single fertilized with high contents of heavy metals. The maximum
ovule growing into an inverted seed covered by an altitude is 240 m a.s.l. The buoyant seeds are eas-
epimatium. Seeds including the epimatium obovoid ily transported by running water and germinate in
to pyriform, with a curved beak at inverted apex and mud temporarily falling dry with fluctuating water
a short, usually persisting crest at distal end, 18–20 tables in lakes and streams. The fleshy epimatium
× 12–13 mm, green or glaucous becoming fleshy and is consumed by birds, and perhaps fish, but also of-
deep red when ripe; seed proper subglobose, 15–17 ten rots away, even on the plant. The thick, fibrous
× 10–11 mm, rough and porous (floating in water). bark is perhaps an adaptation against damage from
 stones transported in flash floods that may occur af-
Description
ter torrential downpours over the sparsely vegetated
plain discharge into the rivers. There are few other Trees to 30 m tall, to 80 cm d.b.h., bole straight and
plants that occupy this habitat; near the Chûtes de la erect, clear of branches for 15 m or more in large
Madeleine on the Rivière des Lacs Dacrydium guil­ trees growing in forest. Bark not described. Branches
lauminii occurs at the water edge with R. minus, and spreading, forming a rounded crown. Foliage branch-
there are some sedges (Carex sp.). es alternate or subopposite, spreading but becoming
somewhat pendulous in shaded branches, with a few
dispersed, decurrent, spreading, small scale leaves on
Conservation
leading shoots and more numerous and larger, pho-
940 Retrophyllum minus is a rare species with a limited tosynthetically dominant leaves on lateral branch-
distribution and a very narrowly defined habitat. In lets. Leaves on lateral branchlets spirally inserted
extremely dry periods it is vulnerable to dehydra- but pectinately arranged, appearing opposite, with
tion and small subpopulations could be wiped out a decurrent base and a 90° twist at the point where
by brush fires. These fires have increased in fre- the leaf blade abruptly widens and is free from the
quency in recent decades due to increased human shoot, twisting in opposite directions on either side
visits and activity, e.g. pine plantations and geologi- of shoot, orientating blade on one side of shoot with
cal surveys involving road building, on the Plaine adaxial side uppermost and on other side with adax-
des Lacs. Much of this plain falls within a mining ial side downwards. Leaf shape in mature trees (ob)
concession. Currently, this species is only protected ovate to elliptic, free part (5–)8–11 × 2–3.5 mm, widest
in a single small botanical reserve at the Chûtes de la in or just above the middle, more or less abrubtly nar-
Madeleine, which covers only one small subpopula- rowing to an obtuse or weakly apiculate apex; midrib
tion. The total number of mature individual plants narrow and conspicuous on both sides. Stomata on
is unknown, but lies somewhere below 2500 and is both sides (leaves amphistomatic), numerous but not
inferred to be declining because of habitat distur- in regualar lines. Pollen cones not known. Seed cones
bance, destruction and fragmentation. on lateral branchlets with small leaves, subterminal
IUCN: EN [B1ab(iii)+2ab(iii); C2a(i)] fertile bract scale-like, 1–3 × 2 mm, with a single ferti-
lized ovule growing into an inverted seed covered by
an epimatium. Seeds including the epimatium ellip-
Uses
soid, with a prominent thin crest at distal end, 20–22
There are no uses recorded for this species. It is in × 10–14 mm, green becoming fleshy and brownish
cultivation in a few botanic gardens as a pot-grown red when ripe; seed proper 12–14 × 7–8 mm, with a
young plant. small beak at micropylar end.

Taxonomic notes
Retrophyllum piresii (Silba) C. N. Page, Notes Roy.
Bot. Gard. Edinburgh 45: 380. 1989. Decussocarpus This species was first described and named as Dec­
piresii Silba, Phytologia 54: 461. 1983; Nageia piresii ussocarpus piresii from a single collection; its taxo-
(Silba) de Laub., Blumea 32 (1): 211. 1987. Type: nomic base is therefore not very convincing. The
Brazil: Rondonia, Serra dos Pacahas Novos, [in type collection N. A. Rosa & J. M. Pires (et al. on
the National Park], N. A. Rosa & J. M. Pires 856 duplicates in K and MO) 856 (not 586 as published
(holotype US). in the protologue!) has duplicates at K, MO and US
(holotype) and appears to differ from those I have
seen of R. rospigliosii in its smaller, narrower leaves
Etymology
and its more oblong seeds. Other stated differences,
This species was named after one of the collectors, such as leaf shapes, are more difficult to assess for
J. M. Pires, who gathered the type collection. lack of more material. On the other hand, this genus
had not been known from Brazil previously (the only
other South American species, R. rospigliosii (Pilg.)
Vernacular names
C. N. Page, occurs in Colombia, Ecuador, Peru, and
None have been recorded. Venezuela and has a strictly Andean ­distribution).
More material is needed, but as yet lacking, to as- growing in forest. Bark brown, weathering blackish
certain the taxonomic status of this species more se- grey, exfoliating in large flakes. Branches ascend-
curely. It should at present be provisionally accepted ing or spreading, forming a pyramidal and eventu-
as distinct. ally rounded crown. Foliage branches alternate or
subopposite, spreading but becoming pendulous in
shaded branches, with dispersed, decurrent, spread-
Distribution
ing, small scale leaves on leading shoots and more
Brazil: Rondônia (Serra dos Pacahas Novos). numerous and larger, photosynthetically domi-
TDWG codes: 84 BZN-RN nant leaves on lateral branchlets. Leaves on lateral
branchlets spirally inserted but pectinately arranged,
appearing opposite, with a decurrent base and a 90° 941
Ecology
twist at the point where the leaf blade abruptly wid-
No details are known about the habitat of this spe- ens and is free from the shoot, twisting in opposite
cies other than that its type specimen has been col- directions on either side of shoot, orientating blade
lected at an altitude of 250 m. The area appears to on one side of shoot with adaxial side uppermost
be well forested viewed via Google Earth© satellite and on other side with adaxial side downwards. Leaf
imagery (accessed 13 March 2009). shape ovate-lanceolate or ovate-elliptic, longest in
juvenile plants, much shorter in top of old trees, also
shorter at base and end of lateral branchlets, free
Conservation
part (6–)10–18(–25) × 3–5(–6) mm, mostly widest
Only known from Rondonia (Serra dos Pacahas just below the middle, tapering to an acute or apicu-
Novos) and occurring within a national park; new late apex; midrib narrow and conspicuous on abaxial
localities recently reported have not been verified side, inconspicuous on adaxial side. Stomata on both
by examination of herbarium specimens and need sides (leaves amphistomatic), numerous but not in
confirmation. regualar lines. Pollen cones at or near ends of foliage
IUCN: DD branchlets, axillary to leaves or in groups of 3–5 to-
gether with one terminal, globose when immature,
5–7 mm long, 2–2.5 mm diam. at anthesis; micro-
Retrophyllum rospigliosii (Pilg.) C. N. Page, Notes sporophylls in spirals, triangular, with an acuminate
Roy. Bot. Gard. Edinburgh 45: 380. 1989. Podo­ apex, each bearing two pollen sacs. Seed cones on
carpus rospigliosii Pilg., Notizbl. Bot. Gart. Berlin- lateral branchlets with small leaves; subterminal fer-
Dahlem 8: 273. 1923; Decussocarpus rospigliosii tile bract leaf-like, 3 × 2 mm, with a single fertilized
(Pilg.) de Laub., J. Arnold Arbor. 50: 347. 1969; ovule growing into an inverted seed covered by an
Nageia rospigliosii (Pilg.) de Laub., Blumea 32 (1): epimatium. Seeds including the epimatium subglo-
211. 1987. Type: Peru: Pasco, Oxapampa, N. Esposto bose to ovoid-pyriform, with a short crest at distal
556 (holotype USM). end, 18–25(–30) × 13–18(–20) mm, green or glaucous
becoming fleshy and dark red when ripe, drying to a
hard shell; seed proper 13–17 × 9–12 mm, with a nar-
Etymology
row beak at micropylar end.
This species was named after Dr C. J. Rospigliosi of
the Natural History Museum in Lima, Peru.
Distribution
Colombia, Ecuador, Central Peru, W Venezuela
Vernacular names
(Merida and Tachira).
pino hayuelo, pino laso, pino de montana, pino TDWG codes: 82 VEN 83 CLM ECU PER
romeron, romerillo macho (Spanish)
Ecology
Description
Retrophyllum rospigliosii occurs in montane tropical
Trees to 45 m tall, to 1.8 m d.b.h.; bole straight and rainforest, in which it can attain large size. Its altitu-
erect, clear of branches for 15 m or more in large trees dinal range is from 1500 m to 3300 m (–3750 m in Co-
 lombia and Peru) a.s.l., so it occurs in wet rain­forest
Vernacular names
up to high altitude cloud forest or mossy forest. It can
form more or less extensive pure stands on exposed dakua salusalu, kau solo (Viti Levu); mugo (New
sites, but is more often found scattered among angio­ Guinea, Danau Paniai [Wissel] Lakes)
sperms or sometimes with Prumno­pitys spp.
Description
Conservation
Trees to 45(–60?) m tall, to 1.5(–2?) m d.b.h.; large
This timber species is under much pressure from trees with spur-butresses at base; bole straight and
logging and most of the formerly quite extensive erect, terete, clear of branches for 20 m or more in
942 stands have now disappeared or are reduced to large trees growing in forest. Bark smooth, eventu-
a few trees. It is considered still in decline as log- ally with faint vertical fissures, light to dark brown,
ging and forest clearing have left several trees from weathering blackish grey, exfoliating in small dis-
which herbarium collections were taken by David integrating longitudinal flakes. Branches ascending
de Laubenfels in Venezuela standing alone in pas- or spreading, forming a pyramidal and eventu-
ture since 1980. This situation is undoubtedly also ally rounded crown. Foliage branches alternate or
found in the other countries where this tree occurs subopposite, spreading but becoming pendulous
and in Peru populations are becoming reduced in in shaded branches, with remotely dispersed, de-
number of mature trees (see e.g. Reynel et al., 2006). current, appressed, small and thin scale leaves on
IUCN: VU (A 4c-d) leading shoots and more numerous and larger,
photo­synthetically dominant leaves on lateral
branchlets. Leaves on lateral branchlets spirally in-
Uses
serted but pectinately arranged, appearing opposite,
This species is a valuable timber tree and can yield with a decurrent base and a 90° twist at the point
large sizes of sawn timber. Its wood is of very good where the leaf blade abruptly widens and is free
quality, straight grained, of medium density, durable from the shoot, twisting in opposite directions on
and workable. It is much used for construction, car- either side of shoot, orientating blade on one side of
pentry, cabinet making, and wood turning. Mature shoot with adaxial side uppermost and on other side
trees are often left standing from forest transformed with adaxial side downwards. Leaf shape ovate-lan-
into pasture for cattle and serve to give shelter from ceolate or ovate-elliptic, longest in juvenile plants,
sun and heat for the animals. In future they may be much shorter in top of old trees, also shorter at base
planted for the same purpose; to what extent this is and end of lateral branchlets, free part (10–)15–25
already being done is not known. (–40) × 3–5(–7) mm, mostly widest below the mid-
dle, especially juvenile forms gradually tapering to
an acute, or in short leaves obtuse apex; midrib nar-
Retrophyllum vitiense (Seem.) C. N. Page, Notes row and conspicuous on abaxial side, inconspicu-
Roy. Bot. Gard. Edinburgh 45: 380. 1989. Podo­ ous on adaxial side. Stomata on both sides (leaves
carpus vitiensis Seem., Bonplandia 10: 366. 1862; amphistomatic), numerous but not in regualar
Nageia vitiensis (Seem.) Kuntze, Revis. Gen. Pl. 2: lines. Pollen cones on relatively short subterminal
800. 1891; Decussocarpus vitiensis (Seem.) de Laub., or lateral branchlets with scale leaves, in groups
J. Arnold Arbor. 50: 342. 1969. Type: Fiji: Western of 2–3 together with one terminal, ovate when im-
Division, Viti Levu, B. C. Seemann 576 (lectotype K, mature, becoming long cylindrical, 10–25 mm long
sheet 2/2, here designated). and 2–2.5 mm diam. at anthesis; microsporophylls
in ranks of tight spirals, triangular, keeled, with an
Podocarpus filicifolius N. E. Gray, J. Arnold Arbor. acuminate apex, each bearing two pollen sacs. Seed
43: 74. 1962. cones on 6–12 mm long lateral branchlets with scale
leaves; subterminal fertile scale(s) slightly longer
than sterile scale leaves, with a single fertilized ovule
Etymology
growing into an inverted seed covered by an epima-
The species epithet refers to Viti Levu, the island in tium. Seeds including the epimatium more or less
Fiji from where it was first described. pyriform, with a short crest at distal end, 14–20 ×
10–13 mm, green or glaucous becoming fleshy and in montane forest with Serianthes sp., Syzygium sp.,
deep red when ripe; seed proper subglobose, 12–16 Dacrycarpus imbricatus, Podocarpus sp., and scat-
× 8–10 mm, with a narrow beak at micropylar end. tered palms and frequent thickets of bamboo; here
a tree was reported to be 60 m tall. Its altitudinal
range is from near sea level to 1800 m a.s.l. In Fiji
Taxonomic notes
it is a common constituent of lowland to montane
Lectotype collection number designated by De rainforest containing several other conifers: Agathis
Laubenfels in J. Arnold Arbor. 50: 342 (1969). At K macrophylla, Dacrycarpus imbricatus, Dacrydium
there are two sheets of Seemann 576, one (sheet 1/2) nausoriense, D. nidulum, and Podocarpus neriifolius,
is a seedling (complete) and one (sheet 2/2) a branch as well as various angiosperm trees including palms.
from a young tree corresponding with the plate in 943
the protologue. They cannot be both the lectotype as
Conservation
they come from two separate plants. The latter sheet
is to be designated as the lectotype. Despite logging of this and other podocarp trees in
many areas of its wide range, which undoubtedly
has led to some reduction of its area of occupancy
Distribution
(AOO), since logged forest is often converted to
Malesia: Maluku [Moluccas]; Papuasia: New Britain, other land uses that do not bring back these trees,
New Guinea; Southwest Pacific: Santa Cruz Islands, Retrophyllum vitiense is as yet not considered a
Fiji Islands. threatened species.
TDWG codes: 42 MOL 43 BIS NWG-IJ NWG-PN 60 IUCN: LC
FIJ SCZ
Uses
Ecology
This species is an important and valuable timber
Retrophyllum vitiense is a tall tree of tropical low- tree which can reach large size. The wood is traded
land to montane rainforest, usually occurring as within Fiji and used for heavy exterior construction
scattered individual emergents. In New Guinea it is timber, house building (e.g. floors), and furniture.
sometimes frequent in forests dominated by Agathis Uses elsewhere within its natural range are similar.
labillardierei and Lithocarpus spp. In the Bismarck This tropical tree is in cultivation only in some bo-
Archipelago (New Ireland) it was found growing tanic gardens.
 Saxegothaea Lindl., J. Hort. Soc. London 6: 258. 1851. [& Paxton’s Flower Gard.
2: 111. 1851] (nom. cons.). [“Saxe-Gothaea”]. Type: Saxegothaea conspicua Lindl.
(Podocarpaceae).

Squamataxus J. Nelson, Pinaceae: 168. 1866 (nom. il­ mm; base petiolate, twisted; blade with a thin mid-
leg.). Type: Squamataxus albertiana J. Nelson [Saxe­ rib on both sides, at upperside yellowish green or
gothaea conspicua Lindl.]. more often lustrous dark green; flushing leaves
pinkish red or yellowish green; apex mucronate.
Named after Prince Albert of Saxe-Coburg-Gotha Stomata in two centrally placed, whitish bands on
944 (1819–1861), Prince Consort to Queen Victoria and abaxial (lower) side, arranged in continuous lines.
a zealous supporter of science and the arts. Pollen cones axillary, situated on branchlets below
seed cones or separately, solitary or sometimes in
pairs, subtended by 2–3 bracts, cylindrical, 4–6 mm
Description
long, 1.5 mm wide at anthesis; microsporophylls spi-
See the species description. rally arranged, numerous, minute, with two pollen
sacs. Seed cones terminal on short, axillary branch-
lets with deciduous scale leaves, globular with pro-
Distribution
truding scale tips at maturity, 9–12 mm diam., light
As for the species. glaucous green, purplish and fleshy when ripe. Cone
scales 15–20, spirally arranged, at first imbricate;
proximal part of (mostly) fertile scales swelling on
Saxegothaea conspicua Lindl., J. Hort. Soc. London both sides so that the tips stand out. Seeds 1 per scale
6: 258. 1851. Squamataxus albertiana J. Nelson, (from 2 inverted ovules), enclosed, subglobose, ca.
Pinaceae: 168. 1866 (nom. illeg.). Type: Chile: 3 mm diam., slightly flattened, lustrous tan or yel-
[“Patagonia”], W. Lobb 81 (holotype not located, lowish with a dull hilum.
isotype K). Fig. 315, 316
Distribution
Etymology
S Argentina: Chubut, Neuquén, Rio Negro; S Chile:
The species epithet means ‘remarkable’ or ‘striking’ Aisén, Biobio, La Araucania, Los Lagos, Maule.
(English conspicuous from Latin conspicuus). TDWG codes: 85 AGS-CB AGS-NE AGS-RN CLC-BI
CLC-LA CLC-MA CLS-AI CLS-LL
Vernacular names
Ecology
Prince Albert yew (England); manio, maniu, manio
macho, manio hembra (Chile) Saxegothaea conspicua occurs in wet (Valdivian)
rain forest, usually along streams in the Andean part
of its range, but on slopes of mountains in very high
Description
rainfall areas in the coastal mountains. Its altitudinal
Monoecious evergreen trees to 25(–30?) m tall; range is from near sea-level (in the coastal ranges) to
trunk 1–2 m diam., monopodial or with multiple ca. 1000 m a.s.l. in the Andes. North of 38° S it be-
stems. Bark smooth, exfoliating in rounded or ir- comes scarce due to the transition to a drier climate.
regular flakes, brown turning purplish, weathering It is an extremely shade tolerant tree that can grow
blackish grey only where flakes persist. Branches under canopy of (evergreen) angiosperm forest,
long, ascending and spreading, eventually droop- reminiscent of Taxus in the northern hemisphere.
ing or pendulous, forming a broad, rounded crown. This species is often associated with Podocarpus
Leaves spirally arranged, pectinate, short decur- nubigenus, Laureliopsis philippiana, Dasyphyllum
rent, spreading in two divergent planes on shaded diacnathoides, Embothrium coccineum, Notho­fagus
branchlets, more irregularly in outer, sun-exposed obliqua, N. nervosa, N. dombeyi, Weinmannia tricho­
foliage, linear or falcate, (10–)15–25(–30) × 2–3.5 sperma, and trees and shrubs of Myrtaceae. Most of
these trees and shrubs are evergreen, as is Saxegoth­ IUCN: NT
aea conspicua, with the exception of Nothofagus spp.
Little is known about seed dispersal, it is assumed
Uses
that birds will be involved.
The wood of this species is used when straight for
contstruction work and carpentry; more often trees
Conservation
occur that do not grow straight boles and these are
Despite its wide distribution logging still threatens put to firewood. In general, it is not considered an
this species. Increasing demands for firewood make important timber tree. This species is well estab-
much of this cutting indiscriminate, on the other lished in cultivation as an amenity tree in countries
hand selective logging aims at relatively young, with relatively abundant rainfall and mild winters, 945
straight trees. Conversion of natural forest to planta- where it makes a large, domed and usually multi-
tion forest with exotic tree species and to agriculture stemmed tree when planted free standing in parks
further reduce its extent of occurrence (EOO) and and large gardens. Planted in woodland surrounded
area of occupancy (AOO); if these trends continue by other trees, its habit will be more upright with
it may reach a threatened status. On the other hand a narrow crown. Cultivars are not known from this
it occurs in many protected areas both in Argentina species.
and Chile.
 Sciadopitys Siebold & Zucc., Fl. Japon. 2 (1): 1. 1842. Type: Sciadopitys verticillata
(Thunb.) Siebold & Zucc. [Taxus verticillata Thunb.] (Sciadopityaceae).

Greek: skias = canopy, umbel; pitys = pine or fir thin chartaceous brown scales. Leaves much re-
tree; referring to the verticillate, spreading cladodes, duced, scale- or cataphyll-like, 3–5 × 2–4 mm, the
hence ‘umbrella pine’. distal, free portion scarious, yellowish green, soon
brown, caducous. Cladodes (phylloid shoots)
mostly on shortened sections of shoots, axillary to
Description
short, triangular scale leaves, arranged helically in
946 See the species description. pseudowhorls of (8–)10–25(–40), spreading, linear,
mostly straight, (3–)8–10(–13) cm long, 2–3.5 mm
wide, with a narrow longitudinal groove from base
Distribution
to apex adaxially and a wider, deeper groove from
As for the species. base to apex abaxially; margins thick, entire, distally
converging to retuse or emarginate apex; surface
smooth, lustrous light to dark green; abaxial groove
Sciadopitys verticillata (Thunb.) Siebold & Zucc., lined with white papillae concealing the stomata,
Fl. Japon. 2 (1): 3, t. 101–102. 1842. Taxus verticillata which are confined to this groove. Pollen cones in
Thunb., in Murray, Linn. Syst. Veg., ed. 14: 895. terminal or sublateral clusters of 8–20, helically ar-
Mai-Jun 1784. Type: Japan: Honshu, [locality not ranged, effectively forming a cone ca. 2 × 1.5 cm,
stated], C. P. Thunberg UPS 23787 (holotype UPS). each individual cone subglobose or broadly ovoid,
Fig. 317, 318 4–6 × 3–5 mm, yellow turning brown. Microspo-
rophylls helically arranged, imbricate, subpeltate,
near abaxial base bearing two yellow pollen sacs.
Etymology
Seed cones terminal or sublateral, from a large, scaly
The species epithet refers to the whorled phyllotaxis bud, ovoid to cylindrical, truncate, often irregular,
of the leaf-like cladodes. 3.5–11 × 2.5–5 cm, maturing in second year, closed
at first, often exuding white resin, bract portion of
bract-scale complex brown, ovuliferous scale por-
Vernacular names
tion green, entire cone turning brown with spread-
Japanese umbrella pine; Koyamaki (Japanese) ing scales. Bract-scale complexes helically arranged
on a stout axis, more or less dolabriform, up to 3.5
× 3.5 cm; bracts and ovuliferous scales proximally
Description
fused, with ovuliferous scales exceeding the truncate
Trees to 40–45 m tall, evergreen, monoecious; trunk bracts and these terminating in a thick ridge with a
monopodial, columnar, up to 3(–3.5?) m d.b.h. Bark central, ligulate, 2–4 mm long, thin, caducous bract
thin and scaly, becoming fissured on trunk, exfo- tip; outer surface of ovuliferous scale portion deeply
liating in long, thin strips and flakes; inner bark grooved, glabrous; cones when falling often partially
reddish brown; outer bark dull brown. Branches disintegrated. Seeds inverted, 8–13 × 6–8 mm (in-
spreading horizontally or occasionally ascending, cluding 2 wings), compressed; base truncate; apex
forming a pyramidal crown in young trees and a acute; surface smooth, brown with white hilum.
narrowly conical or cylindrical crown in mature
trees. Foliage consisting of cladodes (‘needles’) in
Taxonomic notes
pseudowhorls on shortened terminal sections of
long shoots, the density of this foliage dependent on For a detailed discussion, with references, of the seg-
shoot elongation. Long shoots glabrous, green, soon regation of the Japanese umbrella pine as the sole
turning light brown, grooved longitudinally, di- extant member of a distinct family Sciadopityaceae
morphic, with long sections alternating with much (and why it is not a yew [Taxus] nor a pine [Pinus]),
shortened, thicker sections, growing from terminal, I refer to A Monograph of Cupressaceae and Sciado­
conical, obtuse buds 3–5 × 2–4 mm with ­imbricate, pitys (Farjon, 2005a).
 subsequent conversion to managed or planted for-
Distribution
est, or other land uses, has restricted the occurrence
Japan: Honshu (Aichi, Fukushima, Hiroshima, Kyo- of Sciadopitys in many regions. Most of the remain-
to, Nagano, Nara, Okayama, Wakayama), Shikoku, ing stands in Honshu are now only small remnant
Kyushu (Myazaki). populations and larger stands in more or less undis-
TDWG codes: 38 JAP-HN JAP-KY JAP-SH turbed forest are confined to Shikoku and Kyushu.
As this was the situation already 60 years ago, it is
evident that in the past century Sciadopitys has ex-
Ecology
perienced a decline of at least 30% and probably 50%
Locally common in mixed conifer-angiosperm so that is must be listed as Vulnerable.
forests, with Chamaecyparis obtusa, C. pisifera, IUCN: VU (A1c+2c) 947
Tsuga sieboldii, Abies firma, Pinus parviflora, and
angiosperms such as Aesculus turbinata, Magnolia
Uses
obovata, Acanthopanax spp., Cercidiphyllum japoni­
cum, and Acer rufinerve. In dense forest usually The durable wood of this species is used for con-
only ferns, mosses and liverworts grow under the struction purposes and to a limited extent for boat
trees, but in some localities shade tolerant shrubs building and certain kinds of furniture. The fibrous
and small trees such as Ilex sugeroki and Skimmia bark was formerly used for caulking boats (oakum).
japonica can thrive. Sciadopitys occurs as solitary Its most valuable commercial use is undoubtedly in
trees as well as in small ‘groves’ or more or less pure the horticultural industry, as it is a popular planted
stands, reflecting events of recruitment and subse- tree in Japan and (more limited) in Europe and the
quent successional stages in the forest. This conifer USA. Apparently it was not introduced in China, as
is most commonly found in rocky, cool and moist were a few other Japanese conifers such as Crypto­
ravines and valleys in mountainous areas, at alti- meria japonica. The Dutch, who were the only
tudes between 200 m and 1700 m a.s.l. ­Europeans allowed to trade with Japan, brought it to
Jawa early in the 19th century; it arrived successfully
in England and the Netherlands in 1861. A small
Conservation
number of cultivars with dwarfed growth, variegat-
Being a component species in mixed conifer-­ ed foliage and/or pendulous branches has been de-
angiosperm forests where Chamaecyparis obtusa is veloped but these are rarely planted. Some of these
commonly the dominant species, past logging and seem to scarcely differ from the species.
 Sequoia Endl., Syn. Conif.: 197. 1847 (nom. cons.). Conservation of this generic
name was unnecessary [Straub et al. in Taxon 57 (2): 646 (2008)]. Type: Sequoia
sempervirens (D. Don) Endl. [Taxodium sempervirens D. Don] (Cupressaceae).

Gigantabies J. Nelson, Pinaceae: 77. 1866. (nom. il­ trunk, deeply fissured, fibrous, exfoliating in long,
leg.). According to Straub et al. in Taxon 57 (2): 646 thin strips, reddish brown to grey. Branches spread-
(2008) this name was not validly published (Art. 32, ing, curved down except in top of tree, in old trees
Art. 33.9) but it has been listed as validly published with many reiterations, with numerous spreading
in Index Nominum Genericorum (ING). Type: Gi­ foliage branches forming a conical or pyramidal, in
948 gantabies taxifolia J. Nelson [Sequoia sempervirens large trees more rounded crown. Foliage branches
(D. Don) Endl.]. usually spreading horizontally and flattened, lateral
(ultimate) branchlets 3–12 cm long, covered with
Named after Sequoya (ca. 1760–1843), a Cherokee decurrent leaf bases, green turning reddish brown,
chief who invented an alphabet for his people. semi-deciduous. Leaves alternate, decurrent, on
shaded shoots pectinate by a twist of lower free
part, not twisted but curved on most fertile shoots,
Description
mostly linear, flattened, up to 25 × 3 mm (usually ca.
See the species description. 20 mm long but variable on the same shoot from
base to middle to apex), straight or slightly curved;
margins entire; apex acute or acuminate, primarily
Distribution
hypostomatic, with stomata adaxially in two broad
As for the species. bands separated by a narrow midrib; leaf colour lus-
trous green or dull grey-green to glaucous green, the
stomatal bands whitish or glaucous. Pollen cones on
Sequoia sempervirens (D. Don) Endl., Syn. Conif.: the same branches as seed cones, subterminal or ter-
198. 1847. Taxodium sempervirens D. Don, in minal from buds in leaf axils, solitary, ca. 5 × 4 mm;
Lambert, Descr. Pinus 2: [24], t. 7, f. 1. 1824. Type: microsporophylls 10–15, helically arranged, peltate,
USA: California, Santa Cruz Co., Santa Cruz Mts., with erose-denticulate margins, bearing 2–3 abaxial,
[“California: Santa Cruz Mr. Menzies” on back of globose pollen sacs. Seed cones terminal on short
sheet], A. Menzies s.n. (holotype BM). Fig. 319, 320 branchlets, 15–30 × 13–18 mm, maturing to purplish
brown and finally (reddish) brown. Bract-scale com-
plexes 18–25, helically arranged, parting when cone
Etymology
matures except distal 2–3, peltate; distal part irregu-
The species name means ‘evergreen’ and was given larly diamond-shaped, 6–10 × 4–5 mm, rugose, with
to contrast this conifer with the deciduous Taxo­ a central depression and transverse ridge in which
dium distichum, then thought to belong in the same the bract tip forms a 1.5–2.5 mm long process until
genus. it erodes; proximal part narrowing to a pedicellate
base. Seeds 60–100 per cone, irregularly shaped, 5–6
× 3–4 mm, flattened, with angular margins, reddish
Vernacular names
brown, often with dark spots; wings 2, rudimenta-
Redwood, Coast Redwood, California Redwood ry, forming a 1 mm wide continuous margin. This
species is the only known hexaploid (chromosome
count 2n = 6x = 66) in conifers.
Description
Trees to 100–110(–112) m tall, evergreen, monoecious;
Distribution
trunk monopodial, straight, erect, often strongly
buttressed or with large burls at base, sprouting af- W USA: Pacific Coast region from SW Oregon
ter damage, resulting in multi-stemmed trees, up to (Curry Co.) to California (Monterey Co.).
6–7 m diam. above buttress. Bark 2–30 cm thick on TDWG codes: 73 ORE 76 CAL
 been removed by logging and replaced by other tree
Ecology
species or a different type of land use from forest.
The famous Redwood forests form a very distinctive ‘Old growth’ Redwood forest has been greatly re-
lowland coastal vegetation type dominated by this duced since Europeans arrived and covers currently
tallest of all trees. The altitudinal range is (1–)30– ca. 80,000 ha, but almost all of this is now in pro-
750(–920) m a.s.l. Though pure stands of Sequoia are tected areas. Its easy regeneration both from seeds
not uncommon especially at low altitudes on river and resprouting from stumps indicate the resilience
flats near the coast, mixed stands with both conifers of this tree against (man-made) disturbances. On
and angiosperms are more the rule. Most common the other hand, its late successional to climax domi-
among associated conifers are Pseudo­tsuga men­ nance coupled with shade tolerance means it is eas-
ziesii, Tsuga heterophylla and Abies grandis, most ily replaced by more light demanding conifers such 949
frequent angiosperm trees are Acer macro­phyllum as Pseudotsuga menziesii (Douglas Fir). This can be
and Lithocarpus densiflorus. Especially in hilly ter- made ‘permanent’ if forests are chosen to be so man-
rain of upland sites many other tree species can be aged, as indeed they tend to be in commercial for-
present and Sequoia often becomes a lesser compo- estry operations. It is therefore considered that the
nent of the forest. The autecology of this species is long-term maintenance of the present abundance of
remarkable for its capacity to resprout after damage this species is at least in part dependent on conser-
at all levels up the tree, but especially from ligno- vation action.
tubers forming massive, largely subterranean swell- IUCN: NT
ings. Evidence of fire in old growth Redwood forests
attests to its adaptation to such disturbances mainly
Uses
in this manner; many trees have formed several
large stems around an old burnt stump. Longevity as The wood of this tree has been used extensively in
evidenced from ring counts of boles exceeds 2000 the past for construction work wherever decay re-
years, but may on account of this survival strategy sistance was a requirement. Many of the older houses
be substantially longer. The occurence of frequent in California and Oregon were built with it, includ-
oceanic fog and the interception of this moisture ing the shingles on the roofs. Railroad sleepers and
through branchlet disposition, phyllotaxis and leaf bridge timbers were other uses to which it was put.
shape in the crowns of especially old growth trees With the sharp decline in ‘old growth’ stands of this
are apparently crucial to species survival in this oth- tree, which yielded the best quality saw timber for
erwise summer dry region. many of these purposes, uses have more recently
shifted to specialities like turned wood products,
cabinets and polished table tops from its burls or
Conservation
burrs, which produce knotty wood with beautiful
The conservation issues involving Sequoia semper­ patterns and colours. The thick, fibrous, and tough
virens pertain largely to the necessary preserva- bark is used for insulation or mulched for applica-
tion of the remaining ‘old growth’ Redwood forest tion in gardens to suppress weeds in flower beds. The
for ecological reasons and involve much less ques- Coast Redwood is extensively cultivated as an orna-
tions about survival in the wild of the species. The mental tree in many countries and about 10 cultivars
“redwood belt” as given on published maps of dis- are in the trade. In favourable conditions growth is
tribution comprises forest with a (co)dominance rapid and trees can reach impressive sizes within a
of Sequoia sempervirens, forest with the species as century and, if their longevity in their natural range
a minor constituent, as well as ‘potential’ sites, i.e. is even remotely approached, they could continue to
where the species has grown historically but has grow for many more centuries.
 Sequoiadendron J. T. Buchholz, Amer. J. Bot. 26: 536. 1939. Type: Sequoiadendron
giganteum (Lindl.) J. T. Buchholz [Wellingtonia gigantea Lindl.] (Cupressaceae).

Steinhauera Presl, in Sternberg, Fl. Vorwelt 2: 202,

Description
t. 49, 57. 1838. Type: Steinhauera gigantea (Lindl.)
Kuntze ex Voss [Wellingtonia gigantea Lindl. (Se­ Trees to 90–95 m tall, evergreen, monoecious;
quoiadendron giganteum (Lindl.) J. T. Buchholz)]; trunk monopodial, straight, erect, often strongly
Wellingtonia Lindl., Gard. Chron. 1853: 819, 823. 1853, buttressed or with large root bases or burls at base,
non Meisn. 1840. Type: Wellingtonia gigantea Lindl. becoming very massive, up to 8–10 m diam. above
950 [Sequoiadendron giganteum (Lindl.) J. T. Buchholz]; buttress. Bark eventually 30–60 cm thick on lower
Washingtonia Winslow, Calif. Farmer 2: 58. 1854. part of trunk, deeply fissured, fibrous, soft, slowly
(nom. rej.). Type: Washingtonia californica Wins- exfoliating in small, thin strips, cinnamon to red-
low [Sequoiadendron giganteum (Lindl.) J. T. Buch- dish brown. Branches heavy (to 2–2.5 m diam., in
holz]; Americus Hanford, Great Calif. Tree: 6. 1854 older trees often exceeding the leader), spreading
[nom. rej. prop. (1815) by Straub et al. in Taxon 57 (2): and assurgent, with numerous spreading or droop-
645–647 (2008)]. Type: Americus gigantea (Lindl.) ing foliage branches forming a conical or pyrami-
Hanford [Wellingtonia gigantea Lindl. (Sequoia­ dal, in large trees more rounded crown. Foliage
dendron giganteum (Lindl.) J. T. Buchholz)]. branches spreading or drooping, lax, branching al-
ternately and irregularly, the last 3–4 orders covered
Named after Sequoya (ca. 1760–1843); Greek: den­ with green scale leaves, persistent. Leaves helically
dron = tree; the combination was made to denote arranged in 3 ranks, imbricate, decurrent, on mature
relationship with yet separation from the genus Se­ trees variable with branch size, 1–8(–20) × 1–3.5(–5)
quoia. mm, smallest on ultimate branchlets, straight or
slightly curved, bluntly keeled; margins entire, acute
to apiculate; amphistomatic, with scattered lines of
Description
stomata on abaxial face and two more conspicuous
See the species description. lines with more stomata on adaxial side; colour of
adult leaves lustrous dark green. Pollen cones on the
same branches as seed cones but well above them,
Distribution
terminal (rarely subterminal), solitary, subglobose,
As for the species. 4–6 × 3–5 mm; microsporophylls 15–20, helically ar-
ranged, peltate, bearing 2–3 abaxial, globose pollen
sacs. Seed cones terminal on short branchlets, 30–
Sequoiadendron giganteum (Lindl.) J. T. Buchholz, 70(–95) × 25–50(–65) mm, maturing in 2 seasons to
Amer. J. Bot. 26: 536. 1939. Wellingtonia gigantea green and eventually becoming dark brown. Bract-
Lindl., Gard. Chron. 1853: 819, 823. 1853, nom. il­ scale complexes helically arranged, parting when
leg., Art. 53.1. Type: USA: California, Tulare Co., cone matures except distal 2–3, peltate; distal part
Stanislaus River, [“Headwaters of the Stanislaus and diamond-shaped, 12–30 × 6–12 mm, rugose, thick
San Antonio Rivers”], W. Lobb 436 (holotype not and woody; proximal part narrowing to a pedicel-
located, isotype K). Fig. 321, 322 late base. Seeds ca. 200 per cone, irregularly oval,
5–7 × 3–5 mm including wings, much flattened,
greyish brown; wings 2, marginal, of unequal size,
Etymology
1.5–2 mm wide, yellowish brown.
The species epithet refers to the great size of this tree.
Taxonomic notes
Vernacular names
The discovery of this tree by Europeans was an event
Giant Sequoia, Sequoia, Bigtree, Sierra Redwood; well publicised in the popular press of the time, due
Wellingtonia (United Kingdom) to its sensational size which at first caused disbelief
among more cautious scientists. Although first seen
by Albert Kellogg of San Francisco in 1852, speci- mm, but with high year-to-year variation. Tem-
mens reached John Lindley of London late in 1853, perature in winter is mild, with light frosts but oc-
and he first described it (hurriedly) as Wellingtonia casional extremes, and warm, occasionally hot, in
gigantea in the popular Gardener’s Chronicle (of summer. Sequoiadendron giganteum is well adapted
which he was editor) on Christmas Eve of that year to low-intensity forest fires (extremely thick bark)
(Ornduff in Aune, 1994). Kellogg, however, had an and resists windfall exceptionally well, its wood is
American hero (Washington), not a British one, in rot-resistant. As a result its longevity ranges from
mind for the biggest tree on earth. The furious and 2000–3000(–3200) years.
highly nationalistic pathos that ensued, not only in
the popular press, but even in botany, is now amus-
Conservation
ing to read, especially when we realise that Welling­ 951
tonia was, as later homonym, illegitimate and that Although nearly all groves are on public land, enjoy-
Washingtonia soon became the well-known name of ing various levels of protection, the species is under
a palm genus, causing its use for the Giant Sequoia IUCN criteria considered Vulnerable primarily be-
to be officially rejected. cause of historic rates of decline caused by exploi-
tation. Present problems include fire risks, largely
due to (past) management practices which tended
Distribution
to benefit its coniferous competitors (especially
USA: California, Sierra Nevada (Calaveras, Fresno, Abies) rather than the target species, and which have
Madera, Mariposa, Tulare & Tuolumne Co.). greatly accumulated the fuel load for future fires to
TDWG codes: 76 CAL burn more devastatingly. There is a considerable lit-
erature on the conservation aspects of this species,
for a compilation see Aune (1994).
Ecology
IUCN: VU (A1c+d)
This species is forming ‘groves’ of a few to several
thousand individuals in the Mixed Conifer Forest
Uses
belt on the western slopes of the Sierra Nevada. It is
mixed with other conifers: Abies concolor, A. mag­ Since its discovery by Europeans in the mid-19th cen-
nifica, Calocedrus decurrens, Pinus lambertiana, tury, exploitation during the latter half of that cen-
P. ponderosa, P. jeffreyi, Pseudotsuga menziesii, Taxus tury and into the next was considerable. The trees,
brevifolia, and with fewer broad-leaved trees: Quer­ though of high lumber quality and rot-resistant,
cus kelloggii, Q. chrysolepis, Cornus nuttallii, Alnus often shattered on impact of the giant boles. What
rhombifolia, Salix scoulerana, Acer macrophyllum, wood could be used was put into mainly building
and shrubs: Castanopsis sempervirens, Ceanothus applications, and many larger houses in San Fran-
cordulatus, C. parvifolius, C. integerrimus, Arcto­ cisco and the Bay Area were built of its timber. No
staphylos patula, etc. The relatively narrow altitudi- commercial exploitation of wild groves occurs at
nal belt, (830–)1400–2150(–2700) m a.s.l., and the present, and most of these were protected for their
scattered concentration of groves, which tend to scenic and scientific values many years ago. The gi-
become smaller and further apart going north, indi- ant trees are a major international tourist attraction
cate rather narrow climatic and soil conditions that in California. The Giant Sequoia is also highly re-
are optimal in its natural habitat. Most groves are garded as an ornamental tree in parks and gardens
on granite-based residual and alluvial soils, some of large homes and, being easily propagated from
on glacial outwash, and mildly acidic; best growth seed, is sold by many tree nurseries. Several culti-
is on deep, well-drained sandy loams with available vars have been named and are in the trade. The spe-
ground water, the latter appears to be an important cies also has potential as a managed-forest tree for
limiting factor. The climate is humid, with mostly timber production, but has found few applications
autumn rain and winter snow, and dry summers, in commercial forestry thus far.
with mean annual precipitation between 900–1400
 Sundacarpus (J. T. Buchholz & N. E. Gray) C. N. Page, Notes Roy. Bot. Gard.
Edinburgh 45: 378. 1989. Podocarpus L’Hér. ex Pers. sect. Sundacarpus J. T. Buchholz
& N. E. Gray, J. Arnold Arbor. 29: 57. 1948. Type: Sundacarpus amarus (Blume)
C. N. Page [Podocarpus amarus Blume] (Podocarpaceae).

From ‘Sunda’, a name for the Indonesian archipelago tall buttresses. Bark smooth, dark brown weathering
(minus the Moluccas and New Guinea); Greek kar­ blackish grey, in large trees breaking into numerous
pos = fruit. square plates; inner bark red-brown. Branches of
952 trees in forest well above a clear bole, ascending or
erect, then spreading; foliage branches more or less
Description
pendulous. Foliage buds small, globose, with imbri-
See the species description. cate, rounded scales. Leaves on seedlings and juve-
nile plants distinctly rostrate (abrubtly narrowed to
an elongated apex), 3–6(–9) times as long as wide.
Distribution
Adult type leaves usually narrower, broadly linear,
As for the species. 5–15 cm long, 6–15 mm wide, petiolate and with a ta-
pering base, with parallel or more or less undulating
margins, spreading pectinately or assugent, straight
Sundacarpus amarus (Blume) C. N. Page, Notes or irregularly curved, with a central groove over
Roy. Bot. Gard. Edinburgh 45: 378. 1989. Podo­ the midvein adaxially and a raised midrib abaxially,
carpus amarus Blume, Enum. Pl. Javae 1: 88. 1827; usually acuminate at apex, dark green on the upper
Nageia amara (Blume) F. Muell., Select Pl., ed. 2: (grooved) side, light green below. Stomata restricted
138. 1876; Stachycarpus amarus (Blume) Gaussen, to abaxial side (leaves hypostomatic), in two broad
Trav. Lab. Forest. Toulouse T. 2, 1 (2, 20): 105. 1974 bands of numerous densely set lines. Pollen cones
(nom. inval., Art. 33.2); Prumnopitys amara (Blume) on stalks axillary to foliage leaves or subtended by
de Laub., Blumea 24 (1): 190. 1978. Type: Indonesia: deciduous scale leaves, sometimes terminal and
Jawa, [locality not stated], leg. ign. s.n. (holotype L). solitary, more often 3–7 on a stalk, short cylindrical
Fig. 323, 324, 325 when immature, elongating to 15–30 mm, 2.5–4 mm
wide; microsporophylls triangular-trullate, usually
keeled, with serrate or entire margins and two longi-
Etymology
tudinally dehiscing pollen sacs. Seed cones solitary
The species epithet ‘amara’ given by Blume under or with 2–5 on leafless pedunculate shoots 1–4 cm
Podocarpus (= amarus) means ‘bitter’ and refers long, borne subterminally on 3–5 mm long scaly
to the taste of the ‘fruit’ i.e. the fleshy epimatium dwarf shoots, with distal scales with abortive ovules
around the seed. spreading out or recurving and not enlarged or
fused, with sterile scales falling off. Growing seeds
covered by a green, then orange to red and finally
Vernacular names
glaucous purple, fleshy but firm epimatium, becom-
Black pine; choopoola (Queensland); numerous ing nearly spherical with a small protruding apex,
other local vernacular names are applied to this tree; 22–30 mm diam. Seed proper globular, ca. 20 mm
a list of many of these is given in Flora Malesiana, diam., smooth with an indistinct ridge. Cotyledons
ser. 1, 10 (3): 387 (1988). Pilger (1903) cites only kime- in 3 fused pairs.
rak for the Indonesian island of Jawa, but this name
seems to apply to several ‘podocarps’.
Distribution
Malesia: from Sumatera and the Philippines to New
Description
Guinea, New Ireland and New Britain (not in Penin-
Dioecious evergreen trees to 60 m tall, with an sular Malaysia and in Borneo only in Sabah); Aus-
erect trunk to 1.5 m diam., large trees with 3–5 m tralia: NE coastal Queensland.
TDWG codes: 42 BOR-SB JAW LSI MOL PHI SUL stituent of ‘complex notophyll vineforest’, a type of
SUM 43 BIS NWG-IJ NWG-PN 50 QLD-QU tropical rainforest with abundant lianas, where it oc-
curs with Podocarpus dispermus.
Ecology
Conservation
Sundacarpus amarus is a tree of tropical evergreen
rainforests, holding out in primary forest as scat- IUCN: LC
tered tall and sometimes giant emergents and re-
establishing in secondary forest. It is rare at sea
Uses
level, but becomes common from 500 to 2200 m
a.s.l. and has been found to 3000 m on Mt. Kinabalu Sundacarpus amarus is a timber tree of sometimes 953
in Sabah. At lower to middle elevations it is associ- very large dimensions and is logged often together
ated with conifers such as Agathis spp., Dacry­carpus with other podocarps. It is sometimes distinguished
spp., Falcatifolium gruezoi, and Dacrydium spp.; in the timber trade as black podocarp. Its wood is
angiosperms are Cryptocaria pomatia, Dysoxylum, sawn and used for construction (e.g. government
Macaranga, Ficus and many other tree species. It buildings in remote mountain areas of Papua New
grows often in latosols derived from andesite, ba- Guinea), carpentry and joinery, as well as furniture
salt, or granite, rarely in sandy soils or in marshes. making. It is not well known in the trade as it is still
In New Guinea it very common in montane forests being confused with other podocarpaceous species
with Castanopsis, Nothofagus, Calophyllum, Pasania, or treated with other species in the genus Prumno­
Syzygium, and the conifer Phyllocladus hypophyllus. pitys (PROSEA, 1993). Outside some collections in
At the highest altitudes it occurs in mossy forest and botanic gardens it is not known in horticulture or as
becomes stunted. In Queensland S. amarus is a con- a planted forestry tree.
 Taiwania Hayata, J. Linn. Soc., Bot. 37: 330. 1906. Type: Taiwania cryptomerioides
Hayata (Cupressaceae).

Named after the island of Taiwan, from where it was ­ imorphic with tree age; juvenile leaves persistent
d
first described. for 30 years or longer (trees ca. 15 m tall), falcate-­
subulate, (5–) 10–24 × 1.5–3.5 mm, widest near base,
with free part spreading, bilaterally flattened, keeled
Description
on both faces, nearly straight or curved forward,
See the species description. acute-pungent, amphistomatic, glaucous green; adult
954 leaves short lanceolate-trullate, 3–6(–7) × 1.2–3 mm;
Distribution margins entire; apex acute to obtuse, incurved, free
or nearly appressed; leaves amphistomatic, lustrous
As for the species.
(dark) green with whitish stomata. Pollen cones in
terminal clusters of (2–)3–5(–7) on branchlets with
Taiwania cryptomerioides Hayata, J. Linn. scale leaves, ovoid-globose, 2–3 mm long; microspo-
Soc., Bot. 37: 330. 1906. Type: Taiwan: Nantou rophylls numerous, helically arranged, peltate, with
Co., Chiayi-Nantou border, Wusungkengshan, (2–)3(–4) abaxial pollen sacs. Seed cones terminal,
[“ad pedem montis Morrison”], N. Konishi s.n. solitary, maturing in one season to ellipsoid to cy-
(lectotype TI). Fig. 326, 327, 328 lindrical brown cones (9–)12–20(–25) × 6–11 mm.
Cone scales (= bracts) gradually transformed from
Taiwania flousiana Gaussen, Trav. Lab. Forest. Tou- scale leaves below, (12–)14–20(–25), broadly obdel-
louse T. 1 (3, 2): 6. 1939. toid to obtrullate, 6–10 × 5–8 mm; proximal portion
Taiwania yunnanensis Koidz., Acta Phytotax. Geo- cuneate, yellowish brown or reddish brown; distal
bot. 11 (2): 138. 1942. portion dull brown; apex mucronate. Seeds 14–30
per cone (1–2 per scale), ovate-oblong, flat, 4 × 2–3
mm (without wings), light brown or tan. Wings 2,
Etymology
partly overlapping, surrounding the seed, thin fila-
The species epithet means ‘resembling Cryptomeria’ mentous, hyaline, 1–2 mm wide.
and refers to the type of leaves prevalent in young
trees.
Taxonomic notes
The very disjunct occurrence in Yunnan and Myan­
Vernacular names
mar [Burma] in the west and Taiwan in the east
Coffin tree, Taiwania, Taiwan cedar; tai wan shan (the locus classicus) seems to have been the primary
(Chinese) reason why botanists have made taxonomic distinc-
tions. A careful and critical comparison of types
and many other specimens revealed no consistently
Description
different morphological characters (Farjon, 2005a).
Trees to 60–65(–70) m tall, evergreen, monoecious; The recently discovered population in Vietnam fits
trunk monopodial, straight, up to 3–4 m diam. in well with material from the earlier known loca-
above buttressed base. Bark on large trees relatively tions; here it was observed that reversal to juvenile
thin, exfoliating in thin strips and chips, becom- type leaves can occur in lower parts of the crown of
ing fissured, reddish brown or brown weathering mature trees. Transition from juvenile ‘cryptomeri-
grey. Branches spreading or assurgent, lower foliage oid’ foliage leaves to adult ‘sequoiadendroid’ leaves
branches pendulous, forming a conical or pyrami- occurs at an advanced age of trees, even in the wild.
dal crown in young trees, in old trees domed or flat-
topped. Foliage branchlets (sub)pendulous, in young
Distribution
trees with long subulate leaves, in older trees covered
with short, appressed scale leaves, persistent. Leaves China: NW Yunnan, SE Xizang [Tibet]; NE Myan-
alternate to helically arranged, short decurrent, mar [Burma]; Taiwan: Nantou District; Vietnam:
Lao Cai, Van Ban District. [Other reported localities but at about 3000 mm in Vietnam. Reports of this
in China are here considered to be based on intro- species from other areas in China (Fu & Jin, 1992;
duced trees]. Flora of China 4, 1999) do not refer to this type of
TDWG codes: 36 CHC-YN CHT 38 TAI 41 MYA VIE extremely wet monsoon forest and it is very unlikely
that Taiwania is indigenous there.
Ecology
Conservation
Taiwania cryptomerioides is a conifer of montane
forests at altitudes from 1750 m to 2900 m a.s.l. In In Yunnan many stands of old growth Taiwania are
Taiwan it grows in the cool temperate coniferous still liable to be exploited; the establishment of more
forest belt with Chamaecyparis obtusa var. formosa­ reserves for this species is urgent. In Taiwan the es- 955
na and C. formosensis as codominant species and tablishment of Yushan National Park in 1984 pro-
more scattered occurrence of Calocedrus formosa­ tected several natural stands of trees, but many had
na, Cunninghamia konishii, Picea morrisonicola, been felled by that time. Plantation forestry using
Pseudotsuga sinensis, Taxus wallichiana, and Tsuga this species is limited in Taiwan and more extensive
chinensis. Angiosperm trees are common but scat- in Guizhou and Hunan in mainland China; it will
tered, e.g. Castanopsis, Quercus and Trochodendron take many years before these are trees suitable for
aralioides, while shrubs such as Camellia brevistyla, harvest as the species is (beyond sapling and pole
Eurya, Rhododendron, and Vaccinium are more stages) slow growing. In Vietnam, the small popu-
abundant and the bamboo Yushania niitakayamen­ lation is Critically Endangered (CR), because the
sis can cover large areas. In Yunnan and accross the forest remnants in which it still occurs are acutely
border in Myanmar [Burma] it occurs in the mixed threatened by deliberately set fires in the area; it is
coniferous forest with Abies forrestii, Picea brachy­ estimated that perhaps as much as 80% of its habitat
tyla, Larix potaninii, Pseudotsuga sinensis, and Tsuga has already been destroyed. The situation in Myan-
dumosa, and with Cephalotaxus fortunei, Taxus wal­ mar is unknown at present.
lichiana and Torreya grandis var. yunnanensis in the IUCN: VU (A1d)
understorey. All the trees are densely hung with the
lichen Usnea longissima and mosses and leafy liver-
Uses
worts cover trunks and branches. Some angiosperm
trees are mixed in and become more abundant at The wood of Taiwania is very durable and valued
lower altitudes, e.g. Acer, Castanopsis, Lithocarpus, for timber; notable is the (past?) use of it for coffin
Quercus, Magnolia, Schima, and Sorbus, and in the making. In Vietnam it was observed that the local
understorey Rhododendron and many other shrubs H’Mong mountain people use it together with that
are abundant. In Vietnam it is scattered in remnants of Fokienia hodginsii for their houses, in particular
of montane evergreen forest dominated by Fagaceae, for roof planks. Some of this wood is beautifully
Lauraceae and some Magnoliaceae, with only one marked with red and pale yellow annual rings (late
other large conifer, Fokienia hodginsii, common. In and early wood) and prized for furniture. This tree
these forests Taiwania is an emergent, usually form- has been planted in China outside its natural habi-
ing small groves in sheltered side valleys. It can at- tat for much longer than the century that has passed
tain an age of 1600+ (probably over 2000) years and since its botanical description in 1906. It was intro-
belongs ecologically to those conifers which through duced to Japan, Europe, North America, and New
longevity and canopy emergence survive all other Zealand as an ornamental tree, where it grows well
forest trees, ‘waiting’ for episodal forest disturbance outside in temperate regions with none or only light
(most likely fire but possibly also landslides) to re- frosts in winter and sufficient moisture. Despite its
generate. The forest soils are yellow and red acidic attractive form and foliage as a young tree, it has re-
derivatives of granitic or metamorphic rocks. The mained rare and is almost restricted to arboreta and
climate is strongly influenced by monsoons, with botanic gardens.
annual precipitation exceeding 4000 mm in China,
 Taxodium Rich., Ann. Mus. Natl. Hist. Nat. (Paris) 16: 298. 1810. Type: Taxodium
distichum (L.) Rich. [Cupressus disticha L.] (Cupressaceae).

Schubertia Mirb., Nouv. Bull. Soc. Philom. 3: 123.

Distribution
1812 (nom. rej.). Type: Schubertia disticha (L.) Mirb.
[Cupressus disticha L.]; Cuprespinnata J. Nelson, SE North America (USA), Mexico, Guatemala.
Pinaceae: 61. 1866 (nom. illeg.). Type: Cuprespinnata
disticha (L.) J. Nelson [Cupressus disticha L.]
Key to the species of Taxodium
956 The genus Taxus (yew) and Greek: eidos = ‘resem- – Stomata on a leaf most abundant on the abaxial
blance’; referring to the leaves. side, in broader bands than on the adaxial side.
Seed cones mostly 20–35 mm long
T. distichum
Description
– Stomata about equally abundant on both sides
Deciduous or semi-deciduous trees, monoecious; of the leaf. Seed cones mostly 14–25 mm long
trunk monopodial, erect; base often swollen, T. mucronatum
fluted or buttressed, ‘pneumatophores’ or ‘knees’
present or absent. Resin canals in leaves. Bark fis-
sured, exfoliating in more or less fibrous, long strips. Taxodium distichum (L.) Rich., Ann. Mus. Natl.
Branches spreading or ascending, branching sym- Hist. Nat. (Paris) 16: 298. 1810.
podially, forming a domed or flat-topped crown in
large trees. Foliage branches dimorphic, with long
Etymology
and short shoots; long shoots indeterminate; short
shoots determinate, alternate, distichously spread- The species epithet describes the distichous or two-
ing from long shoots, or assurgent to nearly erect. ranked position of the leaves on foliage shoots.
Winter buds prominent, 3–5 mm, adventitious buds
present. Leaves helically arranged, leaf bases short
Vernacular names
decurrent, the proximal part of the free blade twist-
ed or not, if so spreading the leaves in two ranks, if Bald-cypress
not in 5–8 ranks, linear or (short) acicular, amphis-
tomatic. Pollen cones arranged in spike-like to pan-
Description
iculate systems on clustered (sympodial) branchlets,
solitary or in pairs; microsporophylls helically ar- Trees deciduous, to 40(–46) m tall, to 2–3.5(–5) m
ranged, bearing abaxially in two rows near lower d.b.h.; trunk monopodial, erect; base often swollen,
margin 2–10 pollen sacs. Seed cones terminal and fluted or buttressed; ‘pneumatophores’ or ‘knees’
often clustered, near end of last year’s long shoots. common, numerous on root systems of trees grow-
Cones subglobose or more or less ovoid but irregu- ing in or near water, conical, 1(–4) m tall. Bark on
lar 12–40 mm long. Bract-scale complexes ca. 20–30, trunk fissured, exfoliating in more or less fibrous,
helically arranged, partly fused at base and/or apex long strips, light brown, turning grey. Branches
of cone (infertile scales), others opening slightly, spreading or ascending, forming a conical or py-
breaking away, thick woody, peltate, of unequal ramidal crown in young trees (or in cultivation), old
size and irregularly shaped; bracts entirely includ- trees becoming increasingly flat-topped with long
ed or with a minute recurved tip emerging. Seeds main branches in upper half of tree. Foliage short
15–40 per cone, triangular or angular-ovate, oblique, shoots distichously spreading from long shoots, or
more or less flattened, wings very small or vestigial. assurgent to nearly erect, mostly 8–10 cm long, de-
Cotyledons 3–9, usually 4–6, juvenile leaves at first ciduous. Winter buds prominent, 3–5 mm, globose,
4-whorled, soon alternate, similar to adult leaves. with 6–8 ovate, carinate, acute scales. Leaves on long
shoots deciduous, on short shoots mostly persistent,
2 species. spreading in two ranks, or in 5–8 ranks, linear or
(short) acicular, nearly straight or curved (sub­ulate) the stem. The most commonly associated trees are
or lanceolate; free part 3–15(–22) mm long, ca. 1 mm Nyssa aquatica and N. sylvatica var. biflora, which
wide; margins entire; apex obtuse, mucronate, acute form with Taxodium distichum a distinct vegetation
or pungent; stomata in two broad bands separated type. Other common associated angiosperm trees
by a narrow ridge on abaxial (lower) side, in two nar- are Acer rubrum in the northern part of the range of
rower bands separated by a groove or ridge on adax- T. distichum, and Magnolia grandiflora in the south-
ial side; leaf colour light green, or glaucous green, ern States; Pinus spp., Fraxinus spp., Quercus spp.,
with glaucous green stomatal bands. Pollen cones Liquidambar styraciflua, and shrubs e.g. Ilex spp.,
solitary or in pairs, globose to ovoid-oblong, 3–5 × and Viburnum spp., are also often present. Especial-
2–3 mm, yellowish green turning purplish to brown; ly in the southern States all trees are festooned with
microsporophylls (5–)6–10(–15), ovate-peltate, a trailing bromelioid epiphyte: Tillandsia usneoides. 957
keeled, bearing (2–)4–9(–10) globose pollen sacs. Flooding is frequent or sometimes nearly perma-
Seed cones often clustered, subglobose or more or nent with water levels up to 3 m deep; in riparian
less ovoid but irregular, (15–)20–35(–40) mm long. habitat silt is deposited annually. Away from rivers,
Bract-scale complexes distally rounded, more or clay, muck, or peat are common and in soils with
less four-sided, smooth or more commonly rugose, a high organic content trees grow slower and stay
with a transverse ridge, narrowing to a pedicellate smaller. Salinity above ca. 1% will cause trees to die,
base; external surface glaucous green, maturing to so the species avoids areas flooded by seawater. The
light ochraceous brown or grey-brown; internal sur- climate varies greatly along the extensive range, with
face yellowish brown, often with red resin. Seeds ca. dry and cold winters in the interior along the middle
20–40 per cone, 4–7 mm long, lustrous brown, with section of the Mississippi River and a subtropical,
a large, (sub)lateral, pale coloured hilum; wings very warm and humid climate in S Florida.
small or vestigial, forming up to 3 narrow ridges or
parts thereof ca. 1 mm wide.
Uses
The wood of Taxodium distichum is soft, straight-
Distribution
grained and extremely rot resistant and therefore
SE USA: Alabama, Arkansas, Delaware, Florida, widely used in construction and building of houses,
Georgia, Illinois, Indiana, Kentucky, Louisiana, boats, river pilings and sidings, as well as shingles,
Maryland, Mississippi, Missouri, North Carolina, flooring, garden furniture, greenhouses, cooperage,
Oklahoma, South Carolina, Tennessee, S Texas and fencing and other uses for which durability is desir-
Virginia. able. Outside its natural range it is widely used as an
TDWG codes: 74 ILL MSO OKL 75 INI 77 TEX 78 ornamental tree and it has been introduced as early
ALA ARK DEL FLA GEO KTY LOU MRY MSI NCA SCA as 1640 in England. It is currently being planted on a
TEN VRG large scale as an amenity tree in China. Only a lim-
ited number of cultivars has been selected; this spe-
cies is almost uniformly planted with either of the
Ecology
two varieties here recognized. Planted on the shores
Taxodium distichum is a winter deciduous tall tree of lakes and ponds, this tree often develops 1 m tall
dominant in lowland river flood plains and swamps, ‘knees’ from its root system along the water’s edge,
mostly below 30 m but up to 530 m a.s.l., where it adding to the interest of this species. It grows well
can form extensive forests of nearly pure stands on on higher ground away from water, where ‘knees’
(seasonally) inundated fluvial sediment. The two will not develop. In its native riparian habitat it is
varieties are largely sympatric, but var. imbricarium increasingly recognized as a keystone species in
seems to avoid riparian habitat and to prefer stag- swamp forest ecosystems providing both food and
nant pools and swamps. In permanently wet habitat nesting opportunities for rare birds and other wild-
with anaerobic soil conditions this species develops life and as a natural regulator of floods.
characteristic ‘knees’, which grow upwards from
shallow roots, often at a considerable distance from 2 varieties are recognized:
 Taxodium distichum (L.) Rich. var. distichum.
Description
Cupressus disticha L., Sp. Pl. 2: 1003. 1753. Type:
USA: [locality not known; “Cupressus (disticha) ‘Pneumatophores’ or ‘knees’ uncommon, if present
virginiana”], leg. ign. [Herb. Clifford] HSC 449.2 more rounded (not conical). Short shoots assurgent
(lectotype BM). Fig. 329, 330 or erect. Leaves 5–8 ranked, largely appressed but
with free apices, not twisted at base, short acicular
Cupressus disticha L. var. nutans Aiton, Hort. Kew. 3: (subulate) to narrowly lanceolate, 3–10 mm long,
372. 1789; Taxodium distichum (L.) Rich. subsp. nu­ keeled, acute or pungent.
tans (Aiton) E. Murray, Kalmia 12: 25. 1982.
Distribution
958
Vernacular names
SE USA: Coastal Plain from Virginia to Louisiana,
Bald-cypress, Cypress, Southern Cypress, Swamp Cy- generally not as far inland as T. distichum var. dis­
press, Red-cypress, Yellow-cypress, White-­cypress, tichum.
Gulf Cypress TDWG codes: 78 ALA FLA GEO LOU MSI NCA SCA
VRG
Description
Ecology
‘Pneumatophores’ or ‘knees’ common and numer-
ous, conical, 1(–4) m tall. Short shoots spreading, As for the species, but more often in swamps and
but with occasionally assurgent or nearly erect near lakes more distant from rivers.
branchlets with short subulate leaves on the same
tree, or with intermediate forms. Linear, flattened
Conservation
leaves pectinately arranged, twisted near base, 10–
15(–22) mm long, straight or slightly curved, obtuse IUCN: LC
or mucronate.

Taxodium mucronatum Ten., Ann. Sci. Nat. Bot.,

Distribution
sér. 3, 19: 355. 1853. Taxodium distichum (L.) Rich.
As for the species. var. mucronatum (Ten.) A. Henry, in Elwes &
Henry, Trees Gr. Brit. Ireland 1: 175. 1906. Type:
Mexico: México, Valle de México, Netzahualcoyotl,
Conservation
J. G. Hawkes & J. P Hjerting 1389 (neotype K). Fig. 331
IUCN: LC
Etymology
Taxodium distichum (L.) Rich. var. imbricarium
(Nutt.) Sarg., Sylva North America 10: 152. 1896. The species epithet refers to the apex of the leaves,
Cupressus disticha L. var. imbricaria Nutt., Gen. which may terminate in a small, sharp point or
N. Amer. Pl. 2: 224. 1818. Type: USA: [“Found mucron.
from Florida to North Carolina, in swamps and
ponds more remote from the sea.”] “C. disticha *
Vernacular names
imbricaria”, T. Nuttall s.n. (spec. 1) (holotype PH).
Montezuma Bald Cypress, Mexican cypress; Ahue-
Taxodium ascendens Brongn., Ann. Sci. Nat. (Paris) huete, Ciprés, Sabino (Mexico)
30: 182. 1833; Taxodium distichum (L.) Rich. var. as­
cendens (Brongn.) Mast., J. Hort. Soc. London 14:
Description
248. 1892 [“adscendens”].
Trees (semi-) evergreen, to 30(–43) m tall, to 3–8
m d.b.h.; trunk monopodial or forked to multi-
Vernacular names
stemmed, often branching low above ground; ‘pneu-
Pond cypress, Cypress, Black-cypress matophores’ or ‘knees’ rare on root systems of trees
growing in stream beds, rounded. Bark on trunk along upland rivers and lesser streams. It has, how-
deeply fissured, forming anastomosing ridges, ex- ever, been introduced and planted in many localities
foliating in more or less fibrous, long strips, light away from these water sources since pre-Hispanic
brown to reddish brown, turning grey. Branches times throughout Mexico and Guatemala, but these
spreading or ascending, forming a conical or py- grounds all have (or had) water tables reaching up
ramidal crown in young trees, old trees often forked to the relatively deep root system of the trees. Most
and with long main branches, forming a rounded or of the largest trees are found among these planted
irregular crown. Foliage short shoots distichously specimens. Its altitudinal range is (20–)160–2700 m
spreading from long shoots, mostly 6–16 cm long, a.s.l. Germination requires wet soil conditions, ei-
often drooping, usually deciduous shortly before ther submerged or exposed to air. There is no for-
new foliage emerges. Winter buds globose, with mation of ‘knees’ but some trees have a wide, flat, 959
6–8 ovate, carinate, acute scales. Leaves on long mostly subterranean base suggesting a lignotuber
shoots deciduous, on short shoots mostly persist- (this needs investigation). As a result, T. mucro­
ent, spreading in two ranks, linear, nearly straight natum is a major constituent of gallery woodland or
or slightly curved, the free part 3–15(–20) mm long, forest bordering rivers, arroyos and canyon bottoms
ca. 1 mm wide; margins entire; apex obtuse, mucro- with more or less perennial water at least near the
nate, or acute; stomata in two broad bands on both surface. In many streambeds above-ground water
leaf faces; leaf colour light green, or glaucous green, occurs only episodically for a short time after heavy
with glaucous green stomatal bands. Pollen cones rains, carrying large amounts of sediment. Com-
solitary or rarely in pairs, globose to ovoid-oblong, monly associated tree genera are Platanus, Populus
2–4(–5) × 2–3 mm; microsporophylls 6–10(–15), pel- and Salix, at lower altitudes also Ficus and Inga, while
tate, keeled, bearing abaxially in two rows near the many other taxa can occasionally occur, especially
lower margin (2–)4–6(–9) globose pollen sacs. Seed in the southern part of its range. In the northern,
cones subglobose or more or less ovoid but irregu- more arid part of its range T. mucronatum with its
lar, (12–)14–25(–30) mm long. Bract-scale complex- semi-evergreen foliage (only dropped under mois-
es distally rounded, more or less four-sided, smooth ture stress) forms green ribbons along ‘dry’ stream
or more commonly rugose, often pulverulent, with beds in an often semi-arid landscape. This species is
a transverse ridge, narrowing to a pedicellate base; frost sensitive, unlike T. distichum, a species which
external surface glaucous green, maturing to light can withstand extremely low temperatures.
ochraceous brown or nearly grey; internal surface
yellowish brown, often with red resin. Seeds ca. 15–
Conservation
30 per cone, 4–6 mm long, lustrous brown, with a
large, pale coloured hilum; wings very small or ves- A recent assessment of the conservation status of
tigial, forming up to 3 narrow ridges or parts thereof this species in Mexico (Alanis Flores & Favela Lara,
less than 1 mm wide. 2002) concluded that it is not threatened with ex-
tinction. In Texas, the species is listed as endangered
for the State; in Guatemala its status is unknown.
Distribution
IUCN: LC
Guatemala; Mexico; USA: S Texas, scattered along
the Rio Grande.
Uses
TDWG codes: 77 TEX 79 MXC-DF MXC-ME MXC-
MO MXC-PU MXC-TL MXE-AG MXE-CO MXE-CU This tree has undoubtedly been used as a source of
MXE-DU MXE-GU MXE-HI MXE-NL MXE-QU MXE- timber from time immemorial, but perhaps its best
SL MXE-TA MXE-ZA MXG-VC MXN-SI MXN-SO known pre-Columbian use was as a ‘rainmaking’
MXS-CL MXS-GR MXS-JA MXS-MI MXS-NA MXS-OA tree which also had medicinal properties ascribed
MXT-CI 80 GUA to it. The Aztecs planted it in avenues, gardens, and
plazas of the Valle de México; some of these fea-
tures still exist today (Chapultepec, El Contador).
Ecology
In Mexico, this tradition is now continued, with an-
This species differs ecologically from T. distichum in cient trees being preserved and new ones planted.
several ways. Its natural habitat is strictly riparian Martínez (1950) relates some traditional medicinal
 uses; in some parts of Mexico the foliage is used to
decorate church altars during religious ceremonies.
Its commercial value as a timber tree is limited due
to the softness and weakness of the wood.

960
 figure 304. Pseudo­
tsuga japonica foliage
and young seed cone

fig ure 30 6. Pseudo­tsuga

menziesii tree in Mt. Rainier
N.P., Washington, USA

figure 307. Pseudo­tsuga

menziesii giant trunk in
Olympic N.P., Washington

figure 305. Pseudo­tsuga japonica

seed cones

figure 308. Pseudo­tsuga menziesii

var. menziesii seed cones
figure 309. Pseudotsuga sinensis
var. sinensis trees in Taroko N.P., Taiwan

figure 310. Pseudotsuga sinensis

var. sinensis fallen seed cones
 f i gu re 3 13 . Retrophyllum minus in Rivière des Lacs, New Caledonia figure 311. ­Retrophyllum
comptonii young tree on
Mt. Panié, New Caledonia

figure 31 4. Retrophyllum minus foliage and

immature seeds

fi g u re 3 1 2. ­Retrophyllum f i g u re 31 7. Sciadopitys verticillata

­comptonii foliage and seed foliage and pollen cone buds
figure 316. Saxegothaea conspicua foliage
(­photo M. Gardner)
with pollen cones and seed cones
figure 315. Saxe­
gothaea conspicua tree
(photo C. N. Page)

figure 318. Sciado­

pitys verticillata seed
cone (­photo C. N. Page)
 figure 322. Sequoiadendron figure 323. Sundacarpus amarus
­giganteum pollen cones and seed cones trunk and bark, Lake Barrine, Queensland

figure 319. ­Sequoia

semper­virens at Bull
Creek, California
(­photo R. Van Pelt)

figure 32 0. Sequoia
sempervirens seed cones

figure 32 4. Sunda­
carpus amarus foliage

figure 32 1. Sequoia­
dendron giganteum
in California, USA
(photo E. Parker)
figure 32 6. Taiwania cryptomerioides in Yunnan, China
(photo D. Long)

f i g u re 32 5 . Sundacarpus amarus
pollen cones
 figure 329. Taxodium distichum swamp
forest in North Carolina, USA

figure 327. Taiwania cryptomerioides

tree (photo P. Thomas)
figure 330. Taxodium distichum
var. distichum foliage and seed cones

figure 32 8. Taiwania
cryptomerioides trunk in Taiwan
figure 333. Taxus
baccata foliage and pol-
len cones

figure 331. Taxo­

dium mucronatum in
Oaxaca, Mexico

figure 332. Taxus

­baccata ancient
tree on ­ruined wall
of ­Waverley Abbey
f i gu re 3 3 4 . Taxus baccata foliage and seeds

figure 335. Taxus brevifolia in the Wenatchee Mts., Washington, USA

f i gu re 3 36 . Taxus brevifolia foliage and

seeds

figure 337. Taxus cuspidata var. cuspidata in Korea (photo Y. S. Kim)

f i gu re 3 3 8 . Taxus cuspidata var. cuspidata

foliage and seeds

fi g u re 3 39 . Tetraclinis articulata seed cones figure 3 40. Thuja koraiensis in Korea

(photo M. Gardner) (photo Y. S. Kim)
 figure 3 43. Thuja plicata foliage and seed
cones

f i gu re 3 4 1 . Thuja ­koraiensis f i g u re 3 4 2. Thuja koraiensis foliage

foliage branch (upperside) branch (underside)

figure 3 45. Thujopsis

dolabrata var. dolabrata
foliage (upperside)
figure 3 46. Thujopsis
­dolabrata var. dolabrata
­foliage (underside)

figure 3 4 4. Thuja
­sutchuenensis in Daba
Shan, China (© Bedgebury
­Pinetum)

figure 3 4 7. Torreya
­californica foliage with
pollen cones

figure 3 4 8. Torreya
californica foliage
with seeds
 f i gu re 3 4 9 . Torreya nucifera foliage
with seeds

figure 350. Tsuga chinensis var. chinensis in the mountains of Taiwan

figure 35 1. Tsuga chinensis

var. chinensis foliage and seed
f i g u re 352 . Tsuga forrestii foliage cones
and seed cones

f i gu re 35 3 . Tsuga heterophylla on a ‘nurse log’ in Olympic N.P., Washington

figure 354. Tsuga mertensiana var. ­mertensiana figure 355. Widdringtonia cedarbergensis
foliage and seed cones seed cones
f i gu re 35 9 . Wollemia nobilis bud and foliage

figure 35 7. Wid­
dringtonia whytei on
Mt. ­Mulanje, Malawi
(photo P. Cribb)

figure 358. ­Wollemia

nobilis trees in ­Wollemi
N.P., Australia
(photo J. Plaza)

figure 356. Widdring­

tonia nodiflora seed cones
figure 361. Xantho­
cyparis nootkatensis
trunk in Olympic
N.P., Washington

figure 360. Wollemia

nobilis foliage and pollen
cones
figure 362. Xantho­
cyparis vietnamensis foliage
Taxus L., Sp. Pl. 2: 1040. 1753. Type: Taxus baccata L. (Taxaceae).

Verataxus J. Nelson, Pinaceae: 168. 1866 (nom. illeg.). logical characters. Consequently, taxonomies have
Type: Taxus communis J. Nelson (nom. illeg.) [Taxus differed widely, from the recognition of a single
baccata L.]. species with subspecies and varieties (Pilger, 1903)
to the recognition of 24 species with 55 varieties in
Taxus is the classical Latin name for yews.
the most recent treatment (Spjut, 2007). The more
commonly accepted names amount to about 10 spe-
Description
cies and a few subspecies or varieties; a similar
Dioecious (rarely monoecious) evergreen shrubs or taxonomy is adopted here, with all of Spjut’s new 969
trees. Resin in arils of seeds. Bark reddish or pur- names synonymized with already established taxa.
plish brown, becoming scaly. Foliage branchlets ir- This genus is perhaps a good example of a group
regularly alternate. Leaves helically inserted, twisted of species for which it could be illuminating to ap-
at the petiolate base so as to become pectinate or ply some more explicit methodology than intuitive
distichous, linear or falcate, flattened, soft coria- taxonomy based on the careful observation of speci-
ceous, flexible, with a prominent midrib and two mens, as has been done so far with the widely diver-
pale greyish green or pale yellow stomatal bands on gent results cited above. Morphometric analysis of
abaxial side, lacking resin canals. Pollen cones axil- leaf characters could test these in theory, but may
lary, solitary, usually with each leaf on a fertile shoot, not find many gaps between what are mostly quan-
forming double rows, globose with densely clus- titative characters. Another (and currently popular)
tered, peltate microsporophylls each with 4–9 radi- approach is of course genetics (DNA), but here the
ally inserted pollen sacs containing spherical pollen. comprehensive sampling of all the dispersed areas in
Seed bearing structures axillary to foliage leaves, which the genus is found to grow naturally becomes
consisting of a short shoot with small scales and an a major obstacle to obtain results that we can have
axillary fertile dwarf shoot bearing a single, termi- any confidence in. Due to the wide use of yews in
nal, erect ovule. Seeds hard with a highly sclerified cultivation and their likely vulnerability to genetic
seed coat, partly surrounded by a cup-shaped, non- introgression, genetically uncontaminated material
adnate, greenish, fleshy aril, swelling and becoming is only guaranteed from natural populations isolated
within a year glutinous and succulent, usually red or from any other named taxon, as well as from plants
sometimes orange or yellow; the only non-toxic part in cultivation. There appear to be no easy ways out
of the plant. Seedlings with 2–3 cotyledons. of this dilemma (Stuessy, 2009). A ‘consensus’ tax-
onomy might be the only sensible solution, but that
10 species.
would require taxonomists to agree with each other.
Distribution
Key to the species of Taxus
North America: E Canada and E USA south to Flori-
da; W North America from Queen Charlotte Islands The morphological characters that distinguish be-
(BC) to California, E as far as W Montana. E & S tween species of Taxus are notorious for their lack of
Mexico into Central America to Honduras. Eurasia: variation, often resulting in similar or overlapping
W Europe (including Madeira) to S Scandinavia, states. Species, as recognized here, can often only be
NW Russia and Caucasus; Hindu Kush, Himalaya, determined using a combination of unrelated char-
NE India, Myanmar [Burma]; China, Korea, Japan, acters and observations of sufficient material to ob-
Taiwan, Russian Far East (Primorye); Indochina: serve the range of character states that may occur in
Thailand, Laos, Vietnam. North Africa: Atlas Moun- each. In cultivation, forms may be seen that differ
tains. Malesia: Sumatera, Sulawesi, Philippines. from those that occur in natural populations even
without having been given a cultivar name.
Taxonomic notes
1a. Leaves on lateral branchlets pectinate (spread-
The genus Taxus is considered notoriously difficult ing more or less in a plane); midrib raised only
in terms of delimitation of species using morpho-
 in proximal half on adaxial (upper) side and 9a. Leaves on lateral branchlets linear, with revo-
raised to apex on abaxial side 2 lute margins and a cuspidate or sometimes
1b. Leaves on lateral branchlets usually distichous mucronate apex; papillae on abaxial (lower)
(spreading sideways but not in a plane); midrib midrib few, or absent T. mairei
raised to apex on adaxial side and flat on abax- 9b. Leaves on lateral branchlets linear to lanceolate,
ial side 5 with flat or only slightly revolute margins and a
2a. Papillae present along midrib on abaxial (low- mucronate apex; papillae absent T. wallichiana
er) side of leaves; leaves (1.5–)2–3(–3.5) cm
long T. globosa
2b. Papillae absent; leaves (0.8–)1–2.5(–2.9) cm Taxus baccata L., Sp. Pl. 2: 1040. 1753. Type:
970 long 3 [locality not stated (“Habitat in Europa, Canada”)],
3a. Leaves on lateral branchlets slightly S-curved or Herb. Clifford 464 (lectotype BM). Fig. 332, 333, 334
sometimes straight, (1–)1.5–2.5(–2.9) cm long.
Pollen cones few on a branchlet, forming scant
Etymology
rows T. floridana
3b. Leaves on lateral branchlets straight or curved The species epithet baccata (Latin bacca = berry)
near base, (0.8–)1–2(–2.5) cm long. Pollen means ‘berry-like’.
cones many on a branchlet, forming often
dense rows 4
Vernacular names
4a. Leaves on lateral branchlets 1–2.3 mm wide.
Ripe arils around the seeds 7–9 × 5–7 mm. Low Common yew, European yew; Eibe (German); If
shrubs T. canadensis (French); Tis yagodnyi (Russian)
4b. Leaves on lateral branchlets 1.5–3 mm wide.
Ripe arils around the seeds 9–12 × 7–9 mm.
Description
Shrubs or trees T. brevifolia
5a. Papillae present along midrib on abaxial (low- Shrubs or trees to 20(–28) m tall; trunk of trees
er) side of leaves; leaves flat (in fresh leaves), monopodial, short, or multi-stemmed, frequently
(0.8–)1.5–2(–2.3) cm long T. chinensis fluted and asymmetrical, to 3.5 m d.b.h. but usu-
5b. Papillae absent; leaves (slightly) revolute, rarely ally hollow in large specimens, with reiterated stems
flat, (1–)1.5–4(–4.5) cm long 6 inside the old bole; tree bases often readily coppic-
6a. Leaves on lateral branchlets usually more or ing. Bark thin, exfoliating in large irregular flakes,
less directed upwards, arranged in a V-forma- reddish or purplish brown. Branches numerous, as-
tion (rarely radially arranged), straight or only cending to erect, then spreading or drooping, fre-
curved near base 7 quently reiterating, forming a spreading, rounded
6b. Leaves on lateral branchlets distichous but or pyramidal crown. Foliage branchlets irregularly
not arranged in a V-formation, usually falcate alternate, slender, terete, with fine grooves alongside
(curved along most of their length), sometimes decurrent leaf bases, green turning light brown to
nearly straight 8 grey. Terminal buds small, ovoid, with imbricate,
7a. Leaves on lateral branchlets 1.5–2.5 mm wide, closely appressed, rounded, dark brown scales per-
with a cuspidate apex. Pollen cones 6–8 mm sisting at base of new branchlets; lateral buds fre-
diam. Foliage branchlets yellowish brown to quent, dormant, often causing burrs on stems from
grey T. contorta epicormic new growth. Leaves on lateral shoots
7b. Leaves on lateral branchlets 2.2–3.5 mm wide, more or less distichous, sometimes slightly overlap-
with a mucronate or acute apex. Pollen cones ping, linear, 1–3.5(–4) cm long, twisted at the short
3.5–4.5 mm diam. Foliage branchlets pale or- petiolate base, curved outward near base, falcate to
ange-brown to reddish brown T. cuspidata nearly straight, 2–2.5 mm wide, coriaceous; margins
8a. Leaves on lateral branchlets 2–2.5 mm wide. slightly revolute, more or less gradually tapering to
Pollen cones subglobose, 4–5 mm diam a cuspidate apex. Midrib on adaxial (upper) side
T. baccata raised, ca. 0.3 mm wide, nearly continuous to apex,
8b. Leaves on lateral branchlets (1.5–)2–4(–5) mm on abaxial side flat, ca. 0.5 mm wide and continuous
wide. Pollen cones ovoid, 5–6 × 3–4 mm 9 to apex; leaf colour lustrous dark green above, two
pale green bands below. Stomata in two bands on mixed broad-leaved-conifer forests, again mostly
abaxial side, densely and randomly distributed. Pol- on limestone substrates, and often occupying rocky
len cones axillary, solitary, forming rows on either cliffs and slopes. On acid soils yews perform less well
side of fertile shoots, subglobose, 4–5 mm diam., under canopy and usually do not develop beyond a
short pedunculate with dry, yellowish brown, in- sapling stage in woods. Its northern limits in Scan-
creasingly larger bracts at base; axis slightly elon- dinavia are determined by its sensitivity to severe
gating at anthesis. Microsporophylls 6–15, peltate, frost. Its toxicity (all parts except the red arils around
each with 5–9 partly fused pollen sacs on the un- the seeds) prevent browsing by cattle and sheep, but
derside, creamy white or pale yellow. Seed-bearing not by rabbits and deer, as these animals have devel-
structures axillary, solitary or sometimes in pairs, oped a level of immunity to the dangerous alkaloids.
distributed on underside of foliage branchlets, ses- Apart from seed germination (dispersed by birds), 971
sile, with tiny triangular scales covering a very small Taxus baccata readily regenerates from stumps and
dwarf shoot and a single terminal ovule except the roots (suckers); ancient hollow trees may rejuvenate
micropyle. Aril green at first, covering lower half of constantly in this way. When planted, e.g. in church
seed, swelling to succulent red (rarely yellow) and yards and cemeteries, soil pH seems unimportant;
overtopping the seed, leaving its apex free, cup-like, some of the largest and presumably oldest specimen
9–12 mm long, 7–9 mm wide. Seeds ovoid, slightly trees in NW Europe, in particular Brittany (France)
flattened, rarely 3–4-sided, with a small mucro, 6–7 and the British Isles, are known from such locations
× 4–5 mm, green turning brown or black. and were planted probably since Celtic times.

Distribution Conservation
Europe; North Africa (Atlas Mts.); Macaronesia IUCN: LC
(Madeira); Caucasus; Western Asia: from Turkey to
N Iran.
Uses
TDWG codes: 10 DEN FIN GRB IRE NOR SWE 11
AUT-AU AUT-LI BGM-BE BGM-LU CZE-CZ CZE-SK In the Middle Ages the wood of yew was very much
GER HUN NET POL SWI 12 BAL COR FRA-CI FRA-FR in demand for long-bows and cross-bows and was
FRA-MO POR SAR SPA-AN SPA-SP 13 ALB BUL GRC exported from Switzerland to England. Yews were
ITA-IT ITA-SM KRI ROM SIC-SI TUE YUG-BH YUG- also planted near sacred wells, early Christian
CR YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14 churches, monasteries, and castles for symbolic/
BLR BLT-ES BLT-LA BLT-LI KRY RUC RUE RUN RUS religious reasons (Bevan-Jones, 2002) as well as
RUW UKR-MO UKR-UK 20 ALG MOR-MO 21 MDR 33 practical (military) ones. It still is one of the obliga-
NCS TCS 34 IRN TUR tory cemetery trees in NW and Central Europe. The
hard, slow growing wood is used for gates, furniture,
parquet floors, panelling, and is excellent for carving
Ecology
and wood turning as its contorted growth and ‘burls’
Taxus baccata is capable of growing under (not en- form intricate, vari-coloured patterns. For the same
tirely closed) canopy of beech (Fagus spp.) as well as reasons yew does not provide timber suitable for
other deciduous broad-leaved trees, but it will only construction. The toxicity to cattle and horses has
develop to large trees in more open situations. In led to extermination of Taxus baccata from many
Switzerland, the richest area of Central Europe for woodlands in past centuries, when almost all wood-
yew, it forms a yew-beech wood on cool, steep marl land served for grazing animals. Although of lower
slopes in the Jura and the foothills of the Alps up to concentration than in some other species, its alka-
1400 m a.s.l. (Ellenberg, 1988). Under the evergreen loid taxanes, contained mostly in the leaves, yield a
yew, nothing else will grow. In England, Taxus bac­ semi-synthesised anti-cancer drug similar to Taxol®
cata is best developed on chalk downs ‒ again on and yew hedge clippings can still be sold to pharma-
steep slopes ‒ and can form extensive stands outside ceutical companies. As an ornamental shrub or tree
the beech woods invading down grassland. In much it reappeared in the formal gardens of the Baroque
of Europe where the climate is less oceanic it sur- period, as it lends itself to clipped hedges and topi-
vives better in mixed forests, coniferous as well as ary of all shapes. This horticultural interest has in
 turn led to the development of numerous cultivars, two bands on abaxial side; papillae present along the
some of which have bright yellow arils around the midrib. Pollen cones axillary, solitary, forming rows
seeds. on either side of fertile shoots, ovoid to subglobose,
2–4 mm diam., short pedunculate with dry, yellow,
increasingly larger bracts at base; axis slightly elon-
Taxus brevifolia Nutt., N. Amer. Sylva 3: 86, t. 108. gating at anthesis. Microsporophylls 5–10, peltate,
1849. Taxus baccata L. subsp. brevifolia (Nutt.) each with 4–6 partly fused pollen sacs on underside,
Pilg., in Engler, Pflanzenr. IV.5 [18]: 113. 1903. Type: creamy white or pale yellow. Seed-bearing struc-
USA: Oregon, [“Columbia woods”], T. Nuttall s.n. tures axillary, solitary, distributed on underside of
(lectotype BM). Fig. 335, 336 foliage branchlets, with small triangular scales cov-
972 ering a small dwarf shoot 2–8 mm long and a single
Taxus brevifolia Nutt. var. reptaneta Spjut, J. Bot. terminal ovule except the micropyle. Aril green at
Res. Inst. Texas 1 (1): 219. 2007. first, covering lower half of seed, swelling to suc-
Taxus brevifolia Nutt. var. polychaeta Spjut, J. Bot. culent orange or red (rarely yellow) and when fully
Res. Inst. Texas 1 (1): 217. 2007. developed overtopping seed, leaving its apex free,
cup-like, 9–12 mm long, 7–9 mm wide. Seeds ovoid,
more or less 2–4-sided, with a small mucro, 5–8 ×
Etymology
4–5 mm, green turning brown or black.
The epithet brevifolia means ‘with short leaves’; the
leaves are not much shorter than those of other spe-
Taxonomic notes
cies.
In Flora of North America 2: 425 (1993) Taxus brevi­
folia was described as being a shrub or tree, “usually
Vernacular names
upright, typically with a central stem;…” Apparently
Pacific yew there are shrubs occurring within its geographical
range which do not develop a single stem, but in-
stead stems are more or less decumbent and through
Description
adventitious rooting of stems and branches that
Dioecious shrubs or trees to 15 m tall (rarely to 25 m) touch the ground (layering) form spreading thick-
with monopodial stem but sometimes with low, as- ets of ascending stems and branches. The habitat
cending and layering stems (branches); trunk to of these growth forms is often (but not exclusively)
60(–130) c.m. d.b.h. Bark scaly; outer bark (exfolia- rocky places, avalanche shutes, talus slopes, and old
ting scales) purplish to brown; inner bark reddish to stream beds in forests. This habit therefore seems
reddish purple. Branches often from low on trunk, to be an adaptation to environmental conditions.
spreading or strongly ascending to assurgent, some- Taxus canadensis is a shrub which often layers and
times drooping at ends, forming a wide, irregular T. baccata regularly forms clumps through its layer-
crown or spreading bush from layered stems. Foli- ing lower branches from a centrally growing parent
age branchlets slender, terete-angular with decur- tree; this is a trait present in the genus as a whole,
rent leaf bases, yellowish to reddish orange, turning even if rarely found in some species. Taxus brevifolia
reddish brown, with persistent bud scales at base of var. reptaneta Spjut, distinguished on the basis of a
each season’s growth. Leaves pectinately arranged, decumbent habit, is therefore better regarded as an
spreading at 60–90° from shoot axis, mostly par- ecotype, not a taxon.
allel but some overlapping slightly, (0.8–)1–2(–2.5)
cm long, linear, straight or curved near short peti-
Distribution
olate base, 1.5–3 mm wide, more or less concave
or with revolute margins; apex mucronate; texture Pacific Coast Region of North America: S Alaska,
coriaceous; leaf colour lustrous light or dark green British Columbia, Alberta; NW USA (California,
adaxially, pale green abaxially; midrib raised es- Idaho, W Montana, Oregon, Washington).
pecially in proximal half of leaf on adaxial (upper) TDWG codes: 70 ASK 71 BRC 73 IDA MNT ORE WAS
side, continuously raised on abaxial side. Stomata in 76 CAL
 Ecology Uses
Taxus brevifolia is a small understorey tree in the As with most yew wood, its application is only for
conifer forests of the cool and wet Pacific coast of special purposes such as wood carving and turn-
North America roughly between 40° and 60° N and ing (mostly done in Japan), bows for archery, gun
on the western slopes of the northern Rocky Moun- stocks, tool handles, small furniture and musical
tains. The altitudinal range of this species is from instruments. It is hard and rot resistant and can
near sea level to 2200 m a.s.l. In a few places in the be used for fence posts. Native tribes of the Pacif-
southern parts of its range it becomes a dominant ic Northwest used the wood for bows and also for
tree. In rocky, open places like avalanche chutes it wedges to split planks off large trees of Western red
may form layering thickets with decumbent to as- cedar (Thuja plicata). Some of the alcaloid chemi- 973
cending stems. In localities with lower rainfall yews cals of bark and leaves known as taxanes are refined
are confined to streamsides; in the densely forested and processed as an anticancer drug (paclitaxel or
inlets of the Queen Charlotte Islands they only grow Taxol®). Especially the bark of this species contains
on the borders between forest and sea. In most of high concentrations of pure paclitaxel (Hageneder,
the Pacific conifer forest yews are rare or occasional. 2007). After initial failures the pharmaceutical in-
In the Coast Range of southern Oregon and north- dustry has now succeeded in semi-synthesizing this
ern California, T. brevifolia is associated with Quer­ drug from similar chemicals in the leaves of yews.
cus garryana or Fraxinus latifolia and also grows The toxicity of all but the arils around the seeds can
along rivers with Alnus rubra or Populus tricho­ kill certain domesticated animals, but leaves other
carpa. ­Taxus brevifolia is very shade tolerant and species unharmed. Pacific yews are not much used
can hold its own under all but the densest canopies in horticulture and are only found planted in some
of the various evergreen conifers of the region. It is arboreta and botanic gardens.
associated with most of the forest forming conifers
of the Pacific Northwest, in particular with Thuja
plicata, Tsuga heterophylla, Pseudotsuga menziesii, Taxus canadensis Marshall, Arbust. Amer.: 151.
and ­Sequoia sempervirens, indicating its tolerance to 1785. Taxus baccata L. subsp. canadensis (Marshall)
shade. Pilg. in Engler, Pflanzenr. IV.5 [18]: 113. 1903. Type:
USA: Vermont, C. G. Pringle s.n. 1877 (neotype US,
orig. mat. unknown).
Conservation
In the 1980’s and early 1990’s this species, formerly Taxus baccata L. var. minor Michx., Fl. Bor. Amer.
little valued in forestry and actually seen as a prob- 2: 245. 1803; Taxus minor (Michx.) Britton ex Small
lem species in managed stands of timber trees, & Vail, Mem. Torrey Bot. Club 4: 167. 1893. [Mem.
suddenly attained economic value because of its Torrey Bot. Club 5: 19. 1893–94]; Taxus canadensis
chemical taxane, a collective name for chemical Marshall var. minor (Michx.) Spjut, J. Bot. Res. Inst.
compounds contained primarily in the bark of this Texas 1 (1): 274. 2007.
species that turned out to have potent anti-cancer
effects in humans. An uncontrolled rush by people
Etymology
eager to make unexpected money from these small
trees by stripping them of their bark raised concerns The species epithet means ‘from Canada’.
from conservationists. It is unlikely, given its great
range and presence in vast tracts of almost inacces-
Vernacular names
sible forest, that this activity resulted in serious de-
cline, but the species nevertheless made it onto the Canadian yew
CITES Appendix II list. Pharmaceutical research
and successes in synthesizing the active substances
Description
have substantially reduced the stripping of wild
growing trees and the species is not considered to be Usually monoecious shrubs to 2 m tall, often with
in danger of extinction. low, ascending and layering stems. Bark thin, reddish
IUCN: NT to reddish purple. Branches spreading to ­assurgent,
 forming a spreading or prostrate bush. Foliage coniferous forest. It occurs frequently under Fagus
branchlets slender, terete-angular with decurrent grandifolia and Tsuga canadensis, which demon-
leaf bases, yellowish to reddish orange, turning red- strates its tolerance to low light levels during the
dish brown, with persistent bud scales at base of growing season. The shrubs will layer and form
each season’s growth. Leaves pectinately arranged, thickets under the right ecological circumstances,
spreading at 60–90° from shoot axis, mostly par- especially in permanently moist places.
allel but some overlapping slightly, (0.8–)1–2(–2.5)
cm long, linear, straight or curved near short peti-
Conservation
olate base, 1–2.3 mm wide, more or less concave
or with revolute margins; apex mucronate; texture IUCN: LC
974 coriaceous; leaf colour lustrous light or dark green
adaxially, pale green abaxially; midrib raised espe-
Uses
cially in proximal half of leaf on adaxial (upper) side,
continuously raised on abaxial side. Stomata in two Due to its straggling, often prostrate habit Canadian
bands on abaxial side, papillae mostly absent. Pollen yew is rarely used in horticulture and only a hand-
cones axillary, solitary, forming scant rows on either ful of cultivars has been selected. Its wood, although
side of fertile shoots, ovoid to subglobose, 2–3 mm with properties not unsimilar to other species of
diam., short pedunculate with dry, yellow, increas- Taxus, is of little use due to small sizes. Like other
ingly larger bracts at base; axis slightly elongating at species in the genus, some of the alkaloids (taxanes)
anthesis. Microsporophylls 5–8, peltate, each with present in bark and leaves have properties useful in
4–6 partly fused pollen sacs on underside, creamy anti-cancer medicine.
white or pale yellow. Seed-bearing structures axil-
lary, solitary, distributed on underside of foliage
branchlets, with small triangular scales covering a Taxus chinensis (Pilg.) Rehd., J. Arnold Arbor. 1: 51.
small dwarf shoot 1–3 mm long and a single terminal 1919. Taxus baccata L. subsp. cuspidata (Siebold &
ovule except the micropyle. Aril green at first (some- Zucc.) Pilg. var. chinensis Pilg., in Engler, Pflanzenr.
times not developing, the seed then covered at base IV.5 [18]: 112. 1903; Taxus cuspidata Siebold &
with scarious scales), covering lower half of seed, Zucc. var. chinensis (Pilg.) C. K. Schneid., in Silva-
swelling to succulent orange or red (rarely yellow) Tarouca, Uns. Freil.-Nadelhölzer: 276. 1913; Taxus
and when fully developed overtopping seed, leaving wallichiana Zucc. var. chinensis (Pilg.) Florin, Acta
its apex free, cup-like, 7–9 mm long, 5–7 mm wide. Horti Berg. 14 (8): 355. 1948. Type: China: Sichuan,
Seeds ovoid, slightly flattened, with a small mucro, Wushan Xian [“North Wushan”], A. Henry 7155
4–5 × 3–4 mm, green turning brown. (lectotype A).

Distribution Etymology
NE North America: from Labrador and Newfound- The species epithet refers to China.
land westwards to Manitoba and Minnesota, south-
wards to Tennessee.
Vernacular names
TDWG codes: 71 MAN 72 LAB NBR NFL-NE NFL-SP
NSC ONT PEI QUE 74 ILL IOW MIN WIS 75 CNT INI Chinese yew; hong dou shan (Chinese); Thong do
MAI MAS MIC NWH NWJ NWY OHI PEN RHO VER Bac (Vietnamese)
WVA 78 KTY TEN VRG
Description
Ecology
Shrubs or trees to 20 m tall; trunk of trees mono­
Taxus canadensis occurs in forests, bogs, swamps, in podial, to 1(–1.5) m d.b.h. Bark thin, exfoliating in
wet and shady gorges and ravines, on rocky banks strips or irregular flakes, variously coloured, red-
and slopes, at altitudes from near sea level to 1500 dish or purplish brown or grey. Branches numer-
m a.s.l. In forests it is a scattered understorey shrub ous, ascending to erect, then spreading or drooping,
in deciduous broad-leaved forest, mixed forest, and forming a spreading, rounded or pyramidal crown.
Foliage branchlets irregularly alternate, slender,
Distribution
terete, with fine grooves alongside decurrent leaf
bases, green turning yellowish green to ochre or China: Anhui, Chongqing, Fujian, S Gansu, N Gu­
bronze coloured. Terminal buds small, ovoid, with angxi, Guizhou, W Hubei, NE Hunan, S Shaanxi,
imbricate, closely appressed, rounded, dark brown Sichuan, E Yunnan, Zhejiang; N Vietnam.
scales which are early deciduous; lateral buds fre- TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
quent, dormant. Leaves on lateral shoots more or SC CHC-YN CHN-GS CHN-SA CHS-AH CHS-FJ CHS-
less distichous, sometimes slightly overlapping, GX CHS-HN CHS-ZJ 41 VIE
short lanceolate to oblong, (0.8–)1.5–2(–2.3) cm
long, twisted at short petiolate to nearly sessile
Ecology
base, straight or curved near base, 2–3.2 mm wide, 975
thick and coriaceous, convex; margins flat in fresh Taxus chinensis occurs in evergreen and deciduous
leaves, more or less abruptly ending in a mucronate broad-leaved forests, frequently along streams. Its
or sometimes obtuse apex. Midrib on adaxial (up- altitudinal range in China is from 1100 m to 2700 m
per) side raised, 0.2 mm wide, continuous to apex, a.s.l. The undergrowth is often dominated by bam-
on abaxial side flat, densely papillate, 0.3–0.4 mm boo thickets. In Vietnam, it occurs in the northern
wide and continuous to apex; leaf colour lustrous limestone karst formations on steep slopes and on
dark green above, yellowish green with two pale yel- ridges in subtropical to tropical evergreen forest at
lowish bands below. Stomata in two bands on abax- altitudes between 900 m and 1500 m a.s.l. Here it
ial side, densely and randomly distributed. Pollen is associated with the conifers Pseudotsuga sinensis,
cones axillary, solitary, forming rows on either side ­Pinus fenzeliana (syn. P. kwangtungensis), Fokienia
of fertile shoots, ovoid, 5–6 mm long, 3–4 mm wide, hodginsii, Tsuga chinensis, Podocarpus pilgeri, Nageia
short pedunculate with dry, yellowish green to pale fleuryi, and more locally Xanthocyparis vietnamen­
brown, increasingly larger bracts at base; axis slight- sis. There are numerous small-leaved angiosperm
ly elongating at anthesis. Microsporophylls 8–14, trees and shrubs and an abundance of epiphytes due
peltate, each with 4–6(–8) partly fused pollen sacs to high rainfall and frequent fog.
on underside, creamy white or pale yellow. Seed-
bearing structures axillary, solitary or sometimes
Conservation
in pairs, distributed on underside in distal part of
foliage branchlets, sessile, with tiny triangular scales Due to its wide distribution and capacity to regener-
covering a very small dwarf shoot and a single ter- ate from stumps, it is doubtful that logging of larger
minal ovule except the micropyle. Aril green at first, trees for its wood has reduced the extent of occur-
covering lower half of seed, swelling to succulent red rence of this species. Logging in Vietnam may have
or orange (often more or less translucent) and over- had a bigger impact as populations there are small;
topping seed, leaving its apex free, cup-like, 10–13 it may have reduced the species to some extent to
mm long, 7–10 mm wide. Seeds ovoid or obovoid, the less accessible places. Exploitation of its bark and
slightly flattened, with two obtuse ridges and a small foliage for chemical/medical purposes has caused
mucro, 5–8 × 3.5–5 mm, green turning brown or this species to be listed on CITES Appendix II. Har-
black. vesting can only be sustained from plantations, as
indiscriminately stripping trees of bark and foliage
may kill them. In China such plantations have been
Taxonomic notes
established in several places.
This species has been treated as a variety of Taxus IUCN: LC
baccata, T. cuspidata, or T. wallichiana (the latter
in Flora of China 4: 90–91, 1999) or as a species, as
Uses
I have accepted it here. Its most striking distinction
are the short, thick, papillate leaves, quite different The wood of this species is used in China in con-
from the longer, thin, and flexible leaves of T. walli­ struction, cooperage, for the making of furniture,
chiana. The classifications with T. baccata and T. cus­ and for wood carving and turning. In Vietnam it
pidata reflect much broader species concepts held in is also used for irrigation paddles in rice fields. Ex-
the past by some European botanists. tracts of many parts of the plant (roots, wood, bark
 and leaves) are used in traditional Chinese medicine, wide, with parallel, revolute margins and a cuspidate
while in modern times the pharmaceutical industry apex. Midrib on adaxial (upper) side raised, ca. 0.2
became much interested in the anti-tumor proper- mm wide, nearly continuous to apex, on abaxial side
ties of some of its alkaloids (in particular taxanes, flat, ca. 0.4 mm wide and continuous to apex; leaf
drug name: Taxol®), which appear to be present in colour lustrous dark green above, two pale yellowish
all species in low but varying concentrations in bark green bands below. Stomata in two bands on abax-
and leaves. The seeds contain oils which are also ex- ial side, densely and randomly distributed. Pollen
tracted, but treatment is required to neutralize the cones axillary, solitary, forming rows on either side
poisonous alkaloids. In horticulture it has been used of fertile shoots, ovoid, 6–8 mm diam., short pedun-
in bonsai and to a limited extent as a garden shrub. culate with dry, yellowish green or brown, increas-
976 It is doubtful whether this species is in cultivation ingly larger bracts at base; axis slightly elongating at
outside China and Vietnam. anthesis. Microsporophylls 6–14, peltate, each with
4–9 partly fused pollen sacs on underside, pinkish
turning brown. Seed-bearing structures axillary,
Taxus contorta Griff., Icon. Pl. Asiat.: t. 376. 1854
solitary, sessile, with tiny triangular scales covering
[It. Notes: 351, No. 116. 1848 (descr.); Not. Pl. Asiat.
a very small dwarf shoot and a single terminal ovule
4: 28. 1854 (nomen)]. Type: Afghanistan: Badakh­
except micropyle. Aril green at first, covering lower
shan, Kafiristan, W. Griffith 5002 (lectotype K).
half of seed, swelling to succulent red and overtop-
Taxus fuana Nan Li & R. R. Mill, Novon 7 (3): 263. ping seed, leaving its apex free, cup-like, 9–12 mm
1997. long, 7–9 mm wide. Seeds oblong, slightly flattened,
obtusely ridged on two sides distally, with a small
mucro, 6–7 × 4–5 mm, green turning brown or black.
Etymology
The species epithet means ‘contorted’, but it is un-
Taxonomic notes
clear what William Griffith had in mind when he
chose this epithet. Taxus fuana Nan Li & R. R. Mill was considered by
its authors to be distinct from T. wallichiana and
T. yunnanensis; the latter species is now commonly
Vernacular names
treated as synonymous with T. wallichiana. Recent
West Himalayan yew; mi ye hong dou shan (Chi- morphometric analysis of the Himalayan yews con-
nese); postil (Kashmir); thuner (Hindi) firmed this separation (Möller et al., 2007). ­Spjut
(2007) concluded that the long forgotten name
Taxus contorta Griff. has to replace T. fuana because
Description
they are the same taxon. In Flora of China 4: 89–90
Shrubs or trees to 15(–20) m tall; trunk of trees mo- (1999), three species are keyed out with characters
nopodial, short, or multi-stemmed, to 3 m d.b.h. but that in part have to be observed from living (or at
usually hollow in large specimens, with reiterated least not herbarium pressed) foliage; two of these
stems inside the old bole; tree bases often readily cop- involve the Himalayan and Sino-Himalayan taxa
picing. Bark thin, exfoliating in large irregular flakes, T. contorta and T. wallichiana. Re-examination by
reddish brown. Branches numerous, ascending to me of the herbarium specimens held at the Herbar-
erect, then spreading, frequently reiterating, forming ium at Kew (K) and the Natural History Museum,
a rounded or pyramidal crown. Foliage branchlets ir- London (BM) under T. wallichiana confirmed the
regularly alternate, slender, terete, with fine grooves separation into two taxa. The (fuzzy) dividing line
alongside decurrent leaf bases, green turning yellow- seems to lie in eastern Nepal, with all specimens col-
ish brown to grey. Terminal buds small, ovoid, with lected west from there assignable to T. contorta.
imbricate, closely appressed, acute, brown scales
persisting at base of new branchlets; lateral buds fre-
Distribution
quent, dormant. Leaves on lateral shoots more or less
arranged in V-formation, spaced well apart, linear, China: SW Xizang [Tibet] (Jilong); Central & W
1.5–4(–4.5) cm long, only slightly twisted at short Nepal; India (Himachal Pradesh, Jammu & Kash-
petiolate base, nearly straight or straight, 1.5–2.5 mm mir, Uttar Pradesh); N Pakistan; Afghanistan.
TDWG codes: 34 AFG 36 CHT 40 NEP PAK WHM-
Vernacular names
HP WHM-JK WHM-UT
Japanese yew; dong bei hong dou shan (Chinese);
Ichi-i, Araragi (Japanese); Raramani (Ainu of
Ecology
Hokkaido)
In Xizang [Tibet] this species occurs in pine forests
or in mixed conifer-angiosperm forests, usually un-
Description
der canopy as a secondary layer shrub or tree to 12 m
tall, at elevations between 2500 and 3100(–3400) m Shrubs or trees to 25 m tall; trunk to 1 m (–1.5 m)
a.s.l. d.b.h., often branching low. Branches long and
spreading or ascending, forming a broadly rounded, 977
usually compact crown. Bark thin, shallowly fis-
Conservation
sured, exfoliating in strips or flakes, reddish brown.
Exploited intensively in recent years by extrac- Foliage branches numerous, horizontally spread-
tion of foliage and bark for its alkaloid chemical ing, erecto-patent or nearly erect, smooth with fine
compounds (taxanes) which yield paclitaxel (drug grooves along the decurrent leaf bases, green turn-
name: Taxol®) and similar chemicals, from which a ing pale orange-brown or reddish brown. Terminal
drug against ovarian and breast cancer in humans is buds ovate, with imbricate but partly free, broadly
produced, this species has come under threat of ex- elliptical or ovate, weakly keeled, green with hyaline
tinction. It was originally described from a limited margins, persistent at base of branchlets and turn-
area in the Tibetan Himalayas (Xizang, China) but is ing brown. Leaves distichously spreading, directed
now understood to have a much wider distribution, more or less horizontally or often upwards (forming
encompassing all of the western range of the species a V-shape) from lateral shoots, or sometimes more
commonly known as T. wallichiana. The latter spe- radially spreading, linear, (1.1–)1.5–3(–3.5) cm long,
cies is listed on CITES Appendix II; this should now straight or curved near base, 2.2–3.5 mm wide; leaf
apply to both species. Due to the wide distribution base petiolate, twisted, decurrent; margins mostly
of Taxus contorta it does not at present (2009) meet parallel, revolute, abruptly narrowing to a mucro-
any of the IUCN criteria for a threatened category. nate or sometimes acute apex. Midrib on adaxial
IUCN: NT (upper) side slightly elevated, 0.2 mm wide and con-
tinuous to apex, nearly flat and 0.3–0.4 mm wide on
abaxial side. Leaf colour lustrous dark green above,
Uses
deep green with two pale yellowish green bands be-
Recent discovery of anti-cancer effective com- low. Stomata on abaxial side in two bands separated
pounds in the alkaloid taxanes of this and other spe- by a midrib and margins, numerous, arranged in ir-
cies of Taxus has led to the exploitation of foliage regular lines. Pollen cones axillary, arranged in rows
and bark which effectively kills the shrub or tree. In on second year branchlets, sessile, globose, 3.5–4.5
India, Nepal, and Pakistan this species is commonly mm diam., at base with imbricate, broadly ovate
known as T. wallichiana; a more detailed account of scales increasing in size, turning from pale yellow
uses is given under that species. (on margins) to pale reddish brown. Microsporo-
phylls 9–15 per cone, peltate, each bearing 5–8 obo-
vate-cuneate, pale yellow pollen sacs. Seed bearing
Taxus cuspidata Siebold & Zucc., Abh. Math.-Phys. structures axillary, solitary, with numerous ovate or
Cl. Königl. Bayer. Akad. Wiss. 4 (3): 232. 1846. broadly ovate scales at base, partly enclosing a sin-
gle terminal ovule. Arils with a few pairs of scales at
base, obovate, 10–13 mm long, 9–11 mm wide, with
Etymology
a round opening showing the seed, succulent, deep
The species epithet refers to the leaves terminating scarlet or purplish red. Seeds broadly ovoid or tri-
in a small cusp. chonous-ovoid, 5–6 mm long, 4–4.5 mm wide, with
 3(–4) obtuse ridges below acute or mucronate apex, i­ ndustrial scale it is used to make pencils. The Ainu
lustrous chestnut brown when ripe. people of Hokkaido and Sakhalin made their bows,
as well as the scabbards of hunting knives, from its
wood. The heartwood yields a brown or red dye. Ex-
Distribution
tracts of many parts of the plant (roots, wood, bark
Russian Far East: Kuril Is., Sakhalin, Primorye; and leaves) are used in traditional Chinese medi-
China: Heilongjiang, Jilin, Liaoning, Shaanxi; North cine (treating diabetes) while in modern times the
Korea; South Korea; Japan. pharma­ceutical industry became much interested in
TDWG codes: 31 KUR PRM SAK 36 CHM-HJ CHM- the anti-cancer properties of its alkaloids (taxanes,
JL CHM-LN CHN-SA 38 JAP-HK JAP-HN JAP-KY drug name: Taxol®), which appear to be present in all
978 JAP-SH KOR-NK KOR-SK species in low but varying concentrations. The seeds
contain oils which are also extracted, but treatment
is required to neutralize the poisonous alkaloids. In
Ecology
horticulture it has been so popular for a long time
Taxus cuspidata occurs sparsely in mixed conifer that the species is now rare in the wild, but it is
and conifer-deciduous broad-leaved forests in low- grown nearly everywhere in the urbanised areas of
land to lower montane altitudes from 100 m to 1600 Japan. Numerous cultivars exist, and the species is
m a.s.l. In NE China it occurs in conifer forest with used in bonsai culture. Japanese yew was introduced
Abies nephrolepis, Picea jezoensis, Pinus ­koraiensis, to Europe (England) in 1855 by Robert Fortune and
and Larix gmelinii var. olgensis, in Sakhalin Island more cultivars have been developed here as well as
and northern Japan with Abies sachalinensis, Picea in the USA. A hybrid between T. baccata and T. cus­
glehnii, P. jezoensis, Larix kaempferi, and various pidata (Taxus ×media Rehd.) originated in the USA
angiosperm trees e.g. Acer spp., Betula spp., Popu­ around 1900 and has given rise to further cultivars.
lus maximowiczii, Juglans mandshurica, Sorbus
­aucuparia, Ulmus spp., and Kalopanax ricinifolium. 2 varieties are recognized:
Further south in Japan it is common in the under-
story of woods with Acer spp., Ulmus davidiana var. Taxus cuspidata Siebold & Zucc. var. cuspidata.
japonica, Tilia japonica, Juglans ailanthifolia, Quer­ Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
cus mongolica var. grossesserata, and many other Pilg. in Engler, Pflanzenr. IV.5 [18]: 112. 1903. Type:
species of trees. It grows on a variety of soils derived Japan: [locality not stated], P. F. von Siebold s.n.
from granitic, schistose or serpentine base rocks. (lectotype not designated, syntype K). Fig. 337, 338
The variety nana is mostly found growing on rocky
sea coasts but may also occur on exposed rock out- Taxus baccata L. var. microcarpa Trautv., Mém.
crops in the interior. Acad. Imp. Sci. Saint-Pétersbourg (Sav. Etr.) 9: 259.
1859; Taxus cuspidata Siebold & Zucc. var. micro­
carpa (Trautv.) Kolesn., Vestn. Dal’nevost. Fil. Akad.
Conservation
Nauk SSSR 7: 43, f. 2. 1935; Taxus umbraculifera (Sie-
This species has been listed on CITES Appendix II bold ex Endl.) Lawson var. microcarpa (Trautv.) Sp-
in connection with the exploitation of its foliage for jut, J. Bot. Res. Inst. Texas 1 (1): 279. 2007.
the extraction of chemicals active as a anti-cancer Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
drug. This exploitation was localized and has not Pilg. var. latifolia Pilg., in Engler, Pflanzenr. IV.5
resulted in significant decline throughout the wide [18]: 112. 1903; Taxus cuspidata Siebold & Zucc. var.
range of this species. lati­folia (Pilg.) Nakai, Chôsen Sanrin Kaihô 158:
19. 1938; Taxus caespitosa Nakai var. latifolia (Pilg.)
­Spjut, J. Bot. Res. Inst. Texas 1 (1): 269. 2007.
Uses
Taxus caespitosa Nakai, Chôsen Sanrin Kaihô 158:
The wood of this yew is used in China in construc- 20. 1938; Taxus cuspidata Siebold & Zucc. var. caespi­
tion, cooperage, for the making of furniture and tosa (Nakai) Q. L. Wang, Clavis Pl. Chinae Bor.-Or.,
for wood carving and turning. In Japan it is prized ed. 2: 73. 1995.
for interior finish, household furniture, utensils, Taxus biternata Spjut, J. Bot. Res. Inst. Texas 1 (1):
marquetry, wood turning and sculpture; on a more 266. 2007.
 Taxus floridana Nutt. ex Chapm., Fl. S. United
Description
States: 436. 1860. Taxus baccata L. subsp. floridana
Large shrubs or trees. Leaves 1.5–3.5 cm long, more (Nutt. ex Chapm.) Pilg., in Engler, Pflanzenr. IV.5
or less distichously arranged. [18]: 113. 1903; Taxus canadensis Marshall var. flori­
dana (Nutt. ex Chapm.) Silba, Phytologia Mem. 7:
72. 1984; Taxus globosa Schltdl. var. floridana (Nutt.
Distribution
ex Chapm.) Spjut, J. Bot. Res. Inst. Texas 1 (1):
As for the species. 224. 2007. Type: USA: Florida, [“near Aspalaga”],
H. B. Croom s.n. (lectotype PH).
Conservation
979
Etymology
IUCN: LC
The species epithet refers to its occurrence in ­Florida.
Taxus cuspidata Siebold & Zucc. var. nana hort.
ex Rehd., in Bailey, Cycl. Amer. Hort. 4: 1773. 1902.
Vernacular names
Taxus umbraculifera (Siebold ex Endl.) Lawson
var. nana (Rehd.) Spjut, J. Bot. Res. Inst. Texas 1 (1): Florida yew
281. 2007. Type: Japan: Honshu, Hyogo Pref., Mt.
Hyonosen, G. Murata 44671 (neotype A).
Description
Taxus caespitosa Nakai var. angustifolia Spjut, J. Bot. Dioecious shrubs or small trees to 6 m tall (rarely to
Res. Inst. Texas 1 (1): 268. 2007. 10 m); trunk to 40 c.m. d.b.h. Bark scaly; outer bark
(exfoliating scales) purplish to brown; inner bark
reddish to reddish purple. Branches often from low
Vernacular names
on trunk, spreading or ascending, forming a bushy
Dwarf yew crown, often layering. Foliage branchlets slender,
terete-angular with decurrent leaf bases, yellow-
ish to reddish orange, turning reddish brown, with
Description
persistent bud scales at base of each season’s growth.
Decumbent or low, densely branched shrubs to Leaves pectinately arranged, spreading at (60–)80–
2 m tall. Leaves 1.1–2 cm long, more or less radially 90° from the shoot axis, mostly parallel and spaced
spreading from branchlets. well apart, (1–)1.5–2.5(–2.9) cm long, linear, mostly
slightly S-curved but some straight, 1–2(–2.2) mm
wide, more or less concave or with revolute margins;
Distribution
apex mucronate; texture coriaceous; leaf colour lus-
Japan: W Honshu; Russian Far East: Sakhalin, Pri- trous light or dark green adaxially, pale green abaxi-
morskiy Krai (Bolshoy Pelis). ally; midrib raised especially in proximal half of
TDWG codes: 31 PRM SAK 38 JAP-HN leaf on adaxial (upper) side, continuously raised on
abaxial side. Stomata in two bands on abaxial side,
papillae present along midrib. Pollen cones axillary,
Conservation
solitary, forming scant rows on either side of fertile
IUCN: LC shoots, ovoid to subglobose, 2–3 mm diam., short
pedunculate with dry, yellow, increasingly larger
Uses
bracts at base; axis slightly elongating at anthesis.
The decumbent variety of this species has been in Microsporophylls 5–8, peltate, each with 4–6 partly
cultivation in Europe since 1866, when P. F. von Sie- fused pollen sacs on underside, creamy white or pale
bold introduced it to the Netherlands; it became yellow. Seed-bearing structures axillary, solitary,
known in horticulture as cv. ‘Nana’ when propa- distibuted on underside of foliage branchlets, with
gated from cuttings. The Japanese have undoubtedly small triangular scales covering a small dwarf shoot
had it in cultivation for much longer. 1–2 mm long and a single terminal ovule except the
micropyle. Aril green at first, covering lower half
 of seed, swelling to succulent orange or red (rarely
Conservation
yellow) and when fully developed overtopping the
seed, leaving its apex free, cup-like, 7–10 mm long, This species is considered seriously threatened due
6–8 mm wide. Seeds ovoid, slightly flattened, with to habitat degradation. Artificial lowering of the wa-
a small mucro, 5–6 × 4 mm, green turning brown. ter table due to practises in agriculture is believed
to be responsible for significant water stress within
the plant communities where T. floridana is most
Taxonomic notes
common. Although normally occuring in humid
Spjut (2007) has placed Taxus floridana as a variety forests and thickets and therefore outside the danger
under Taxus globosa, and considered it to occur also of frequent fires that occur in Longleaf-Slash pine
980 in northern Mexico (Nuevo Leon, Tamaulipas, and (Pinus palustris-P. elliottii) woods above the river
Veracruz). If this were accepted, the conservation ravines, the drying of the habitat has increased the
status of T. floridana would change dramatically. risk of fire, to which Taxus is sensitive. Increased
Spjut seems to have based his taxonomic concepts occurrence of fungal attacks may also be connected
in the genus on a few selected micromorphological to these stress factors. The total area of occupancy
characters in mainly the foliage leaves and some- (AOO) is estimated to be less than 10 km2 ; popu-
times bud scales or other characters, without due lation numbers are several thousand mature indi-
consideration of variation within populations and/ viduals in total. However, many are unhealthy and
or analysis of correlation between multiple char- regeneration is poor, usually only observed imme-
acters. There are certainly similarities between the diately near parent trees.
two taxa, such as the slender, slightly curved leaves, IUCN: CR (B1+2c)
but there are also differences, and nothing short of
a comprehensive morphometric analysis of all avail-
Uses
able characters would suffice to establish whether
there were two ‘clouds’ of character states in the data The Florida yew is rare in cultivation and apparently
or only one. These results would then ideally be test- only grown by a few local nurseries in Florida. Due
ed with DNA markers. Pending such research, it is to its suitability as a garden ornamental or hedge
premature to unite these two species into one with shrub and in view of its threatened status, its culti-
two varieties. vation should be recommended.

Distribution
Taxus globosa Schltdl., Linnaea 12: 496. 1838. Type:
SE USA: Florida (along the Apalachicola River in Mexico: Hidalgo, Real del Monte, [“zwischen el
Gadsden and Liberty Co.). canino del paso und Huajalote”], C. A. Ehrenberg
TDWG codes: 78 FLA s.n. (lectotype K).

Ecology Etymology
Taxus floridana is restricted to shaded ravines and The species epithet means ‘like a ball’, which refers to
limestone bluffs along the Appalachicola River in the shape of the (immature) seed.
Florida. These ravines where undisturbed develop
a woodland/forest type dominated by Fagus grandi­
Vernacular names
folia, Magnolia grandiflora and Quercus laurifolia;
other frequent tree components are Ilex spp., Quer­ Mesoamerican yew; Ciprecillo colorado, Pinabete
cus alba, Symplocos tinctoria, and Carya tomentosa. colorado, Ciprés colorado (Spanish).
Sometimes this species occurs in thickets dominant-
ed by Kalmia latifolia; elsewhere it may be associ-
Description
ated with the extremely rare and localized conifer
Torreya taxifolia. These ravine forests are humid and Trees to 15 m tall; trunk to 80 c.m. d.b.h. Bark scaly;
if soils have developed they tend to be slightly acidic outer bark (exfoliating scales) purplish to brown; inner
to neutral sandy loams. bark reddish to reddish purple. Branches ­spreading
or ascending, forming a broadly rounded or irregu- globosa is limited to the highest summits around
lar crown. Foliage branchlets slender, terete-angular 2500–3000 m. Further north in Mexico, in par-
with decurrent leaf bases, yellowish to orange, turn- ticular on the eastern declivity of the Sierra ­Madre
ing brown, with persistent bud scales at base of each Oriental and mountains in Tamaulipas, Oaxaca and
season’s growth. Leaves pectinately arranged, spread- Chiapas, the cloud forest has a more extensive al-
ing at 50–80°(–90°) from shoot axis, mostly parallel titudinal distribution, and there Taxus globosa can
and spaced well apart, (1.5–)2–3(–3.5) cm long, linear, occur in forests dominated by broad-leaved trees,
mostly slightly falcate or S-curved but some straight, especially Quercus spp., Liquidambar styraciflua,
(1.6–)2–2.5(–2.8) mm wide, more or less concave or Magnolia sp., Platanus mexicana, Ulmus mexicana,
with revolute margins; apex mucronate to cuspidate; Carpinus caroliniana, Fagus mexicana, Acer skutchii,
texture coriaceous; leaf colour lustrous (light) green Clethra spp., and the conifers Podocarpus matudae 981
adaxially, pale green abaxially; midrib raised espe- and Pinus strobus var. chiapensis. In Nuevo León it is
cially in proximal half of leaf on adaxial (upper) side, found in the understorey of Abies-Picea forest.
continuously raised on abaxial side. Stomata in two
bands on abaxial side, papillae present along midrib.
Conservation
Pollen cones axillary, solitary, forming scant rows on
either side of fertile shoots, ovoid to subglobose, 3–4 IUCN: NT
mm diam., short pedunculate with dry, yellow, in-
creasingly larger bracts at base; axis slightly elongat-
Uses
ing at anthesis. Microsporophylls 8–12, peltate, each
with (3–)4–5 partly fused pollen sacs on underside, This species is uncommon in cultivation, virtually
creamy white or pale yellow. Seed-bearing structures restricted to botanical gardens and arboreta in re-
axillary, solitary, distributed on underside of foliage gions with mild winters. As with other species of
branchlets, with small triangular scales covering a Taxus, in the 1990’s screening for taxanes (for the
small dwarf shoot 2–4 mm long and a single terminal anti-cancer drug paclitaxel and similar chemicals)
ovule except the micropyle. Aril green at first, cover- was undertaken for this species, and these sub­
ing lower half of the seed, swelling to succulent or- stances when found present were cause of exploita-
ange or red (rarely yellow) and when fully developed tion of this tree for a time.
overtopping seed, leaving its apex free, globose, cup-
like, 10–12 mm long, 9–11 mm wide. Seeds ovoid-
globose, slightly flattened, with a small mucro, 6–7 × Taxus mairei (Lemée & Lév.) S. Y. Hu ex T. S. Liu,
5–6 mm, green turning brown. Illustr. Native & Introd. Lign. Pl. Taiwan 1: 16.
1960. Tsuga mairei Lemée & Lév., Monde Pl. 2 (16):
20. 1914; Taxus chinensis (Pilg.) Rehd. var. mairei
Distribution
(Lemée & Lév.) W. C. Cheng & L. K. Fu, Fl. Hupe­
El Salvador; Honduras; Guatemala; Central, E and hensis 1: 28. 1976; Taxus wallichiana Zucc. var.
S Mexico. mairei (Lemée & Lév.) L. K. Fu & Nan Li, Novon 7
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC- (3): 263. 1997. Type: China: NE Yunnan, Dongchuan
PU MXC-TL MXS-CL MXS-GR MXS-JA MXS-MI MXS- Shi, E. E. Maire s.n. (holotype E).
NA MXS-OA MXT-CI 80 ELS HON GUA
Taxus speciosa Florin, Acta Horti Berg. 14 (8): 382,
t. 6. 1948; Taxus mairei (Lemée & Lév.) S. Y. Hu ex
Ecology
T. S. Liu var. speciosa (Florin) Spjut, J. Bot. Res. Inst.
Taxus globosa is a species occurring in montane to Texas 1 (1): 246. 2007.
high montane cloud forests in the subtropics to near Taxus kingstonii Spjut, J. Bot. Res. Inst. Texas 1 (1):
tropics of Mexico and Central America. Its altitu- 240. 2007.
dinal range is generally between 2000 m and 3000
m a.s.l., but in Tamaulipas, Mexico, a collection was
Etymology
made at only 1380 m a.s.l. In El Salvador and Hon-
duras the cloud forest community with conifers like This species was named after the plant collector
Abies guatemalensis, Cupressus lusitanica and Taxus E. E. Maire.
 Vernacular names Taxonomic notes
Maire’s yew; nan fang hong dou shan (Chinese) Taxus mairei was treated as a variety of T. chinensis
in Flora Reipublicae Popularis Sinicae 7: 443 (Cheng
& Fu, 1978), but transferred to T. wallichiana later
Description
and so treated in Flora of China 4: 91 (1999). It has
Shrubs or trees to 30 m tall; trunk of trees mono­ also been synonymized with T. chinensis, in which
podial, to 1.5 m d.b.h. Bark thin, exfoliating in strips case T. mairei would have priority and the species
or irregular flakes, variously coloured, reddish or should have that name, not T. chinensis. Here I treat
purplish brown or grey. Branches numerous, ascend- them as distinct, although the distinctions are minor
982 ing to erect, then spreading or drooping, forming and confined to leaf morphology only. Spjut (2007),
a spreading, rounded or pyramidal crown. Foliage in a recent taxonomic treatment of the genus, has
branchlets irregularly alternate, slender, terete, with taken the splitting further and segregated T. speciosa
fine grooves alongside decurrent leaf bases, green Florin, commonly treated as a synonym, at varietal
turning orange-brown to purplish brown. Terminal rank from T. mairei var. mairei. This appears to have
buds small, ovoid, with imbricate, closely appressed, been decided mostly on branching patterns, which
rounded, dark brown scales mostly early decidu- in this case I do not consider to provide separable
ous at base of new branchlets; lateral buds frequent, character states, as these are even more prone to en-
dormant. Leaves on lateral shoots more or less disti- vironmental influences during development than
chous, spreading at nearly right angles, linear, 1.5–3.5 leaf shapes and sizes.
cm long, twisted at the short petiolate or nearly ses-
sile base, usually falcate, 2–4 mm wide, thick and
Distribution
coriaceous; margins revolute, more or less gradually
tapering to a cuspidate apex or more abruptly end- China: Anhui, Chongqing, Fujian, S Gansu, N Gu­
ing in a mucronate apex. Midrib on adaxial (upper) angdong, N Guangxi, Guizhou, W Henan, W Hubei,
side raised in a shallow groove, 0.2–0.3 mm wide, Hunan, Jiangsu, Jiangxi, S Shaanxi, Sichuan, Yun-
nearly continuous to apex, on abaxial side flat, non- nan, Zhejiang; Taiwan.
papillate or with few papillae, ca. 0.4 mm wide and TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
continuous to apex; leaf colour lustrous dark green SC CHC-YN CHN-GS CHN-SA CHS-AH CHS-FJ CHS-
above, pale green with two pale yellowish bands be- GX CHS-HE CHS-HN CHS-JS CHS-JX CHS-ZJ 38 TAI
low. Stomata in two bands on abaxial side, in short,
irregular lines and/or randomly distributed. Pollen
Ecology
cones axillary, solitary, forming rows on either side
of fertile shoots, ovoid, 5–6 mm long, 3–4 mm wide, Taxus mairei occurs in coniferous and mixed coni-
short pedunculate with dry, yellowish green to pale fer-angiosperm forests and their more or less open,
brown, increasingly larger bracts at base; axis slight- scrubby margins as a scattered, often understorey
ly elongating at anthesis. Microsporophylls 8–14, tree. It is more common on more or less open slopes
peltate, each with 4–6(–8) partly fused pollen sacs with secondary vegetation after the removal of pri-
on underside, creamy white or pale yellow. Seed- mary forest, on rocky slopes and escarpments, of-
bearing structures axillary, solitary or sometimes ten on limestone. Its altitudinal range is substantial,
in pairs, distributed on underside in distal part of from 100 m to 3500 m a.s.l. In the eastern provin­
foliage branchlets, sessile, with tiny triangular scales ces of China it occurs usually below 1200 m but as
covering a very small dwarf shoot and a single ter- high as 2000 m a.s.l. In Sichuan, Yunnan and also
minal ovule except the micropyle. Aril green at first, in Taiwan it reaches to 3000 m and more. In forests
covering lower half of seed, swelling to succulent red T. mairei often occurs along streams, but in open
or orange (often more or less translucent) and over- terrain it can well grow at a distance from any sur-
topping seed, leaving its apex free, cup-like, 10–13 face water.
mm long, 7–10 mm wide. Seeds ovoid or obovoid,
slightly flattened, with two obtuse ridges and a small
Conservation
mucro, 5–8 × 3.5–5 mm, green turning brown or
black. IUCN: LC
 Uses Vernacular names
The uses of this species are presumably similar to East Himalayan yew; thuner (Hindi) & various
those stated in Flora of China 4: 90 (1999) under other local names; xu mi hong dou shan (Chinese);
Taxus wallichiana and are to provide building mate- tampinur batu (Karo).
rial, to make tools used in agriculture and gardens
in the countryside, furniture, and stationery. It is in
Description
cultivation in China and, as with other species of
Taxus, the leaves have been used to extract taxanes Shrubs or trees to 30 m tall; trunk of trees mono­
for use as an anti-cancer drug. podial, to 1.5 m d.b.h. Bark thin, exfoliating in strips
or irregular flakes, variously coloured, reddish or 983
purplish brown or grey. Branches numerous, as-
Taxus wallichiana Zucc., Abh. Math.-Phys. Cl. cending to erect, then spreading or drooping, fre-
Königl. Bayer. Akad. Wiss. 3: 803, t. 5. 1843. Taxus quently reiterating, forming a spreading, rounded
baccata L. subsp. wallichiana (Zucc.) Pilg., in or pyramidal crown. Foliage branchlets irregularly
Engler, Pflanzenr. IV.5 [18]: 112. 1903. Type: India: alternate, slender, terete, with fine grooves alongside
(eastern), N. Wallich s.n. [ex Herb. Zuccarini] decurrent leaf bases, green turning orange-brown to
(lectotype M). purplish brown. Terminal buds small, ovoid, with
imbricate, closely appressed, rounded, dark brown
Cephalotaxus celebica Warb., Monsunia 1: 194. 1900; scales mostly early deciduous at base of new branch-
Taxus celebica (Warb.) H. L. Li, Woody Fl. Taiwan: lets; lateral buds frequent, dormant. Leaves on later-
34. 1963. al shoots more or less distichous, sometimes slightly
Taxus yunnanensis W. C. Cheng & L. K. Fu, Acta overlapping, linear to lanceolate, (1–)1.5–3.5(–4.5)
Phytotax. Sin. 13 (4): 86. 1975; Taxus wallichiana cm long, twisted at short petiolate or nearly sessile
Zucc. var. yunnanensis (W. C. Cheng & L. K. Fu) base, usually falcate, (1.5–)2–4(–5) mm wide, thin
C. T. Kuan, Fl. Sichuan. 2: 215. 1983; Taxus chinen­ and soft; margins flat or slightly revolute, more or
sis (Pilg.) Rehd. var. yunnanensis (W. C. Cheng & less gradually tapering to a cuspidate apex or more
L. K. Fu) L. K. Fu, Vasc. Pl. Hengduan Mount. 1: 214. abruptly ending in a mucronate apex. Midrib on
1993. adaxial (upper) side raised, 0.2–0.3 mm wide, nearly
Taxus sumatrana (Miq.) de Laub., Kalikasan 7 (2): continuous to apex, on abaxial side flat, ca. 0.5 mm
151. 1978; Cephalotaxus sumatrana Miq., Fl. Ned. wide and continuous to apex, usually without pa-
Ind. [Fl. Ind. Batavae] 2 (7): 1076. 1859. pillae; leaf colour lustrous dark green above, pale
Taxus phytonii Spjut, J. Bot. Res. Inst. Texas 1 (1): 237. green with two pale yellowish bands below. Stomata
2007. in two bands on abaxial side, densely and randomly
Taxus contorta Griff. var. mucronata Spjut, J. Bot. distributed. Pollen cones axillary, solitary, forming
Res. Inst. Texas 1 (1): 260. 2007. rows on either side of fertile shoots, ovoid, 5–6 mm
Taxus florinii Spjut, J. Bot. Res. Inst. Texas 1 (1): 222. long, 3–4 mm wide, short pedunculate with dry,
2007. yellowish green to pale brown, increasingly larger
Taxus suffnessii Spjut, J. Bot. Res. Inst. Texas 1 (1): bracts at base; axis slightly elongating at anthesis.
226. 2007. Microsporophylls 8–14, peltate, each with 4–6(–8)
Taxus obscura Spjut, J. Bot. Res. Inst. Texas 1 (1): 235. partly fused pollen sacs on underside, creamy white
2007. or pale yellow. Seed-bearing structures axillary, soli-
tary or sometimes in pairs, distributed on underside
in distal part of foliage branchlets, sessile, with tiny
Etymology
triangular scales covering a very small dwarf shoot
This species was named in honour of Nathaniel Wal- and a single terminal ovule except the micropyle.
lich (1786–1854), a Danish botanist who amassed the Aril green at first, covering lower half of seed, swell-
famous Wallich Herbarium for the East India Com- ing to succulent red or orange (often more or less
pany, now kept at Kew. translucent) and overtopping seed, leaving its apex
 free, cup-like, 10–13 mm long, 7–10 mm wide. Seeds TDWG codes: 36 CHC-YN CHT 40 EHM-AP EHM-
ovoid or obovoid, slightly flattened, with two obtuse BH EHM-DJ EHM-SI NEP 41 MYA VIE 42 PHI SUL
ridges and a small mucro, 5–8 × 3.5–5 mm, green SUM
turning brown or black.
Ecology
Taxonomic notes
Taxus wallichiana is a small to large understorey
In most of the literature referring to Taxus in the or lower canopy tree in montane, temperate, warm
Himalayas only one species, T. wallichiana, is rec- temperate, and tropical submontane to high mon-
ognized for the entire mountain chain [often re- tane forest, both angiosperm and conifer domi-
984 ferred to as T. baccata subsp. wallichiana (Zucc.) nated, deciduous or evergreen, or in mixed forests.
Pilg.]. More recently, all of the populations occur- In open situations on rocky slopes and cliffs it usu-
ring from central Nepal westward to northern Pa- ally forms a large, broadly spreading shrub. Eleva-
kistan and Afghanistan have been assigned to the tion ranges from 900 m to 3700 m a.s.l. and soils
relatively recently described species T. fuana Nan Li are mostly derived from silicate-bearing rocks, i.e.
& R. R. Mill, which was first known from the Hima- acidic to neutral. Like the European T. baccata it is
layas of SW Xizang [Tibet]. This species has turned easily dispersed by birds and can germinate quickly
out to be synonymous with T. contorta Griff., a much in large numbers on suitable sites. It has a very long
earlier name, which has therefore priority and must life-span (theoretically indestructible) and sprouts
be used instead. It is quite distinct from both T. wal­ from stumps as well as roots. Taxus wallichiana oc-
lichiana and from T. baccata, which does not reach curs in pure stands of limited extent or mixed in
further east than N Iran. Two other species, T. chin­ the understorey of Quercus, Abies, Cedrus deodara,
ensis and T. mairei, were included in T. wallichiana and Picea, or in mixed conifer forest. In Vietnam
as varieties in Flora of China 4 (1999), but here it is T. wallichiana has been found growing in submon-
preferred to treat these as (not very) distinct species. tane evergreen mixed forests associated with the
In Malesia, the species Taxus celebica and T. sumat­ conifers Cephalotaxus mannii, Dacrycarpus imbri­
rana, although considered distinct in recent treat- catus, Keteleeria evelyniana, Nageia wallichiana, and
ments (Farjon, 1998 [2001], several Floras, Spjut Podo­carpus neriifolius. It will form dense thickets on
2007) do not differ consistently in their morpholog- exposed rocky slopes with little tree growth. In the
ical characters from T. wallichiana; indeed, earlier Philippines it occurs on high ridges and mountain
botanists usually identified the specimens from the summits in mossy forest, or sometimes in rocky
Philippines, Sumatera, and Sulawesi as that species. grass and scrubland.
In some of these accounts, T. celebica and T. sumat­
rana are reported to extend from Malesia into In-
Conservation
dochina and China, or even to Nepal, and there to
occur alongside T. wallichiana. Here these two spe- In certain regions of India in particular, harvesting
cies are considered synonymous with T. wallichiana, of leaves and bark for extraction of alkaloid taxanes,
which therefore extends from Nepal to Sulawesi. In used to produce the anti-cancer drug paclitaxel or
this place, it is not possible to comment in detail on similar chemicals, has led to a decline perhaps as
the recent work by Spjut (2007), other than to report much as 90% in recent decades (Molur & Walker,
that comparison of the type specimens of his new 1998). Logging in Yunnan, China may have reduced
species from the region with specimens of T. wal­ the rare populations in that country. It is therefore
lichiana have caused me to conclude that they are prudent to flag this species as NT, even though its
synonyms of that species. extent of occurrence (EEO) is very large and may
include areas where this kind of exploitation has not
occurred. Taxus wallichiana is listed under CITES
Distribution
Appendix II. It occurs in several protected areas, e.g.
China: SE Xizang [Tibet], NW Yunnan; E Nepal; the Sagarmatha National Park in Nepal. Cultivation
Bhutan; India: Arunachal Pradesh, Assam; Myan- on a large scale in the context of pharmacology could
mar [Burma]; Vietnam; Malesia: Philippines, Su- reduce the pressure on wild growing populations.
lawesi, Sumatera. IUCN: NT
 and leaves and sometimes of inner bark for a long
Uses
time in various potions, tinctures, and pastes. The
The wood of Himalayan yew is durable and strong only non-toxic part of yews, the fleshy aril around
and is used for door frames, cabinet work and the seed, is consumed by local inhabitants as jams.
wood turning and wood inlaying, also for candle- The inner bark also produces a red dye, often used in
sticks, knife handles etc. Less refined products are religious ceremonies by Brahmins of Nepal (Singh,
gates and fences, poles, struts and ‘wattle and daub’ 2007). Since the exploitation of Himalayan yew for
in walls of rural buildings. The wood is also burnt its foliage containing taxanes has proved to be un-
as incense in Nepal and parts of Tibet. The leaves sustainable, cultivation efforts are being undertaken
are toxic but can be given as fodder to goats if no in the Himalayan foothills and elsewhere in India;
other foliage is available. The alkaloid compounds this involves both species T. contorta and T. wallichi­ 985
(taxanes) of the bark are a source for the anti-cancer ana. These species, because of their similarity with
drug paclitaxel (Taxol®) which has become a major European yew (T. baccata), are rarely found in culti-
reason for exploitation in recent years. The leaves vation in Europe, but they are used as ornamentals
yield similar chemicals in low concentrations. Tra- elsewhere, as in Baguio, Philippines.
ditional medicine has made use of young shoots
 Tetraclinis Mast., J. Roy. Hort. Soc. London 14: 250. 1892. Type: Tetraclinis articulata
(Vahl) Mast. [Thuja articulata Vahl] (Cupressaceae).

Greek: tetra = four; kline = bed, couch; referring to erals; apical margins denticulate; stomata in two
the leaves in whorls of 4. lateral rows or bands on facials, more sparsely on
laterals; glands subapical, small, most prominent
on facials; abaxial leaf surface smooth, light green.
Description
Pollen cones terminal, solitary, ovoid-globose, ca.
See the species description. 4 × 2.5 mm, reddish, maturing to yellowish brown
986 or brown; microsporophylls 8–12, decussate, peltate,
Distribution obtuse, bearing 4 abaxial pollen sacs. Seed cones sol-
itary or sometimes clustered, terminal on branch-
As for the species.
lets, maturing within one year to (sub)tetragonal
cones 10–13 × 12–17 mm in size and becoming
Tetraclinis articulata (Vahl) Mast., J. Roy. Hort. first glaucous, then purplish brown to light brown.
Soc. London 14: 250. 1892. Thuja articulata Vahl, Bract-scale complexes 4, decussate, nearly equal in
Symb. Bot. 2: 96. 1791. Type: Tunisia: Hammam-el- size, thick woody, ovate-cordate, 10–12 × 8–12 mm;
Lif, [“legi copiose in montibus ad Hamamelis”], leg. lower pair widest; upper pair with a truncate apex;
ign. s.n. A (holotype C). Pl. 41, Fig. 339 all abaxially smooth or rugose, with a subapical,
small, recurved process (bract tip), adaxially striate,
with white seed scars near base, paler than exterior
Etymology
parts. Seeds 4–6 per cone, conical-triangular, 4–5 ×
The species epithet refers to the seemingly articulate 3 mm, brown, with dark spots; basal hilum whitish;
ultimate branchlets. wings 2, at oblique angle, large, ca. 10 × 6 mm, reni-
form, thin menbranous, hyaline.
Vernacular names
Distribution
Sandarac; Thuya d’Algérie, Thuya de Barbarie, Bois
de Citre; Araâr (Morocco); Alerce (Spain) N Algeria, Malta, Morocco, S Spain, N Tunisia, NE
Libya (?); in Malta and Spain only tiny relict popula-
tions remain.
Description
TDWG codes: 12 SPA-SP 13 SIC-MA 20 ALG MOR-
Shrubs or small trees to 6–8(–15) m tall, evergreen, MO TUN
monoecious; trunk monopodial or multistemmed,
usually branching low or forked, up to 50 cm diam,
Ecology
coppicing from base. Bark on large stems becoming
rough, fissured, breaking into small plates, turning Tetraclinis articulata occurs in the semi-arid to
from light brown to grey. Branches long and slender, winter rain parts of the Atlas Mountains and some
more or less crooked angularly, forming a pyramidal scattered localities in the coastal hills near sea level;
crown only in young, undisturbed plants, otherwise there often on limestone, mixed with maquis. More
irregular. Foliage branches articulate, branching al- commonly in subclimax vegetation with Pistacia
ternately at various angles, ultimate branchlets 1–2 lentiscus, Quercus ilex, and Juniperus phoenicea, lo-
mm thick, seemingly costate, persistent. Leaves dec- cally into the zone with Cedrus atlantica, on S-facing,
ussate, appearing in whorls of 4 on thinnest branch- warm and dry slopes to 1800 m, on N-facing slopes
lets, obviously decussate on thicker branches, long to 1300 m a.s.l. Woodland with this species is often
decurrent, adnate to shoot except apex, 1.6–8 × 1–1.5 severely degraded; however, this species survives
mm (still longer on older branches, smallest on ul- both fire and browsing by man’s animals through
timate branchlets), weakly dimorphic; facials linear, its coppicing capacity, unless the pressure becomes
with broader, acute apex, partly covered proximally too severe. Old coppice stools have very dense wood
by the linear-spathulate, transversely convex lat- and may be very ancient, but dating is very difficult.
 5

987

6
4

11

10 9 12

Pl ate 4 1. Tetraclinis articulata. 1. Habit of tree. 2. Branch with foliage and seed cones. 3. Branchlet with
pollen cones. 4. Pollen cone. 5, 6. Microsporophylls with open pollen sacs and pollen. 7. Immature seed
cones. 8, 9. Seed cones. 10. Seed cone scale. 11, 12. Seeds.
 formed burrs (‘lupias’), was prized and used for fur-
Conservation
niture, cabinet making, and wood turning due to its
The populations in Malta and southern Spain are intricate markings and figuring from adventitious
highly threatened. In North Africa, however, the growth. Today, only the larger trunks and stools
estimated area of occupancy is in excess of 1 mil- with their contorted wood structure are valuable;
lion ha (Pardos & Pardos, 1997), with the largest most of the lighter wood is used for fuel locally, es-
populations in Algeria. Despite the capacity to cop- pecially in regions deficient in trees and with a poor
pice, there are indications that increased browsing rural population. The resin (‘sandarac’) is used as in-
pressure, especially by goats, prevent trees from cense in religious ceremonies and also forms a basis
regrowing in many areas (Gaussen, 1968, shows a for varnishes and laquer work; Arabic tribes use(d)
988 photo­graph from Morocco to this effect). it for medicinal purposes. In cultivation it was for-
IUCN: NT merly propagated by the French colonial powers
(Service d’Eau et Forêts) in North Africa in attempts
to extend its use as coppice wood in semi-arid areas.
Uses
It is uncommon as an amenity tree or garden orna-
From the time of Phoenician and Roman coloni- mental and outdoor planting is restricted to coun-
sation in North Africa to the present, the wood of tries with a Mediterranean climate.
Tetraclinis, and especially of the coppiced stools that
Thuja L., Sp. Pl. 2: 1002. 1753. Type: Thuja occidentalis L. (Cupressaceae).

Greek: thyia = resinous tree, also citrus tree. 2a. Seed cones with 6–8(–10) scales. Pollen cones
with 4–6 microsporophylls T. occidentalis
2b. Seed cones with 8–12(–14) scales. Pollen cones
Description
with (6–) 8–16 microsporophylls. 3
Shrubs or small to large trees to 75 m tall, evergreen, 3a. Seed cones with 8–12 scales, the two fertile pairs
monoecious; trunk monopodial or multistemmed; of equal size, more or less truncate. Shrubs or
bark fibrous, exfoliating in longitudinal flakes or small trees T. koraiensis
strips. Branches spreading, curved down or assur- 3b. Seed cones with 8–12(–14) scales, only one pair 989
gent to ascending, forming a conical to pyramidal of equal size, obtuse or acute. Normally large
crown; branching according to Massart’s model. Fo- trees 4
liage branches plagiotropic, branchlets alternate to 4a. Seed cones 10–16(–18) mm long; largest scales
subdistichous, ultimate branchlets often more nu- narrow, l/w ratio > 1.5. Ultimate branchlets on
merous on the acroscopic side of lateral branchlets, foliage sprays more numerous on one (acro-
lateral branchlets compressed, persistent. Leaves scopic) side of lateral branchlets, forming regu-
decussate (semi-whorled), on whip shoots long lar tapering units T. plicata
decurrent, on lateral branchlets dimorphic; facials 4b. Seed cones 7–12(–14) cm long; largest scales
obtrullate-rhombic, smaller than laterals; later- broad, l/w ratio < 1.5. Ultimate branchlets on
als broadly falcate, spreading from apex of facials, foliage sprays alternately arranged, forming ir-
conduplicate; margins entire; most leaves amphisto- regular, more rounded units T. standishii
matic, but upper facials often epistomatic; stomata
most numerous on lower surface of laterals; glands
inconspicuous or absent. Pollen cones terminal, sol- Thuja koraiensis Nakai, Bot. Mag. (Tokyo) 33: 196.
itary; microsporophylls 4–16, each with 2–4 abaxial 1919. Type: North Korea: Hamkyongnam-do Prov.,
pollen sacs. Seed cones terminal, solitary or rarely Samsu-gun, Paek-san, V. L. Komarov 85 (lectotype
in pairs, maturing to ovoid-globose, narrowly ovoid LE). Fig. 340, 341, 342
or elliptical, partly opening cones. Bract-scale com-
plexes 6–12(–14), decussate; lowest pair reduced;
Etymology
2–3 following pairs enlarged and spreading; upper
pairs gradually smaller and narrower and apical pair The species epithet refers to Korea, from where it
often connate. Seeds 4–10 per cone, flattened, with was first known to science.
2 wings. Seedlings with 2 cotyledons.
5 species. Vernacular names
Korean Arbor-vitae ; chao xian ya bai (Chinese);
Distribution
Nioi-nezuko (Japanese)
Asia: disjunct in China: Chongqing (Dabashan), Ji-
lin (Changbai Shan); Korea; Japan. North America:
Description
(disjunct) E Canada & USA; W Canada & USA
(coastal to N California). Shrubs or small trees to 6–10 m in their native habi-
tat; trunk often multistemmed, or monopodial, as-
surgent especially on slopes, to 30 cm d.b.h. Bark
Key to the species of Thuja
on stems thin, exfoliating in longitudinal flakes,
1a. Mature seed cones 5–8 mm long. Foliage sprays reddish brown or purplish brown. Branches spread-
delicate, with very small eglandular leaves ing to assurgent or ascending, forming a low, layer-
T. sutchuenensis ing shrub or a pyramidal crown. Foliage branches
1b. Mature seed cones (7–)8–16(–18) mm long. Fo- plagio­tropic; branchlets alternate to subdistichous;
liage sprays coarse, with small and larger leaves; third year shoots abruptly turning red-brown; lat-
facial leaves usually glandular 2 eral branchlets compressed, with underside more
 flattened than convex upperside, persistent. Leaves of volcanic origin and grows most abundantly on
decussate, imbricate, on lateral branchlets dimor- exposed, granitic slopes and crags with acidic skel-
phic; facials obtrullate-rhombic; laterals broadly etal soil. The climate is cool to cold temperate with
falcate, spreading from apex of facials, conduplicate; high rainfall in the mountains and prolonged winter
leaves variable in size from 1–10 mm long on a single snow cover.
branchlet system (facials smaller than laterals); mar-
gins entire; apex acute to obtuse, of laterals incurved,
Conservation
adnate; stomata most numerous on lower surface of
laterals in a slightly sunken band; glands small, often The extent of occurrence (EOO) calculated for this
visible on facials; leaf colour when newly emerged species, based on distribution data of herbarium
990 often glaucous, turning dark green; stomatal bands collections, is 48,091 km2 and the area of occupancy
white. Pollen cones subglobose, 2–3 × 2–3 mm, pur- (AOO) is unknown but probably less than 500 km2.
ple maturing to purplish black; microsporophylls This small, shrubby tree is not exploited for tim-
6–10, peltate, each with 3–4 abaxial, yellow pollen ber, but may be cut for its foliage used as incense.
sacs. Seed cones solitary or sometimes in pairs, Most subpopulations are small and scattered. The
maturing in one year (2 seasons) to ovoid-globose; only subpopulation within a protected area occurs
opening cones 7–11 × 6–9 mm with brown scales. in China. The herbarium collections consulted from
Bract-scale complexes 8–12, decussate; lowest pair Korea (Farjon, 2005a) are all more than 80 years old.
reduced, recurved; two following pairs enlarged to Rather than interpreting this as a sign of decline,
5–6 × 2–6 mm and spreading; upper pairs gradually I think this is a result of geopolitics affecting bo-
smaller and narrower, to 1 mm wide, often connate; tanical research and distribution of its results to the
larger scales abaxially striate or with longitudinal world’s herbaria. It was not possible consult Korean
grooves and a small, subapical bract tip, adaxially herbaria. The Conifer Specialist Group of IUCN
rugose, with two seed depressions. Seeds 8–10 per wishes to flag this species as near threatened.
cone, ellipsoid, 4–5 × 1–3 mm, flattened, light brown; IUCN: NT
wings 2, marginal, 1–2 mm wide, meeting with a
notch at both ends, thin menbranous, with minute
Uses
puberulence near the distal part.
Because most plants in the wild are shrubs, this spe-
cies is not useful for its wood. It was introduced to
Distribution
the USA by Ernest Wilson in 1917, from where it
NE China: Jilin (Changbai Shan); North Korea came to Europe shortly after; however it remains
(Pyong­annam-do, Pyonganpuk-do, Hamkyongnam-­ relatively rare in cultivation and is mostly restricted
do, Kangwon-do); South Korea (Kyongsanpuk-do, to arboreta and botanic gardens. This is partly due
Kangwon-do, Cholla-namdo). to difficulties of access of new material, as most
TDWG codes: 36 CHM-JL 38 KOR-NK KOR-SK populations occur in North Korea. Remarkably, in
cultivation in the UK, under conditions with ample
moisture as in Hergest Croft Gardens, Hereford-
Ecology
shire, some plants have grown into a tree much
Thuja koraiensis occurs on middle and upper slopes taller than any known in the wild in Korea; it shows
of mountains, at altitudes between 750 m and 1950 no signs of being near its limits at ca. 15 m tall. This
m a.s.l. Where slopes are exposed, it forms dense, form is apparently based on E. H. Wilson’s collection
low thickets, but in sheltered ravines it can become No. 9244 made on 12 October 1917 on Kongo-san at
a small upright tree. It is often associated with Pinus 1500 m a.s.l. in South Korea. In the Netherlands, a
pumila, growing underneath its canopy; on lower very glaucous tree form is grown in the Gimborn
slopes it occurs as a small tree in taller conifer for- Arboretum and University of Utrecht Botanic Gar-
ests with Abies nephrolepis, Picea koraiensis, P. jez­ den. In NE China and Korea the shrubby form is
oensis, Pinus koraiensis, or P. sibirica; less often it used as an ornamental in gardens and some public
grows in clearings in mixed deciduous woodland in parks on a limited scale.
areas with thin, rocky soil. It appears to avoid rocks
Thuja occidentalis L., Sp. Pl. 2: 1002. 1753. Type: or fused; two largest pairs spreading, ca. 8 × 4 mm,
USA: [locality unknown], P. Kalm LINN 1136.1 (ob)ovate-oblong, thin, obtuse or acute, with a sub-
(lectotype LINN). apical bract tip 0.2 mm, abaxially cinnamon or light
brown, dull, adaxially light orange-brown, lustrous,
without apparent seed marks. Seeds 4–8 per cone,
Etymology
usually only 4 (well) developed, 4 × 1–2 mm, flat-
The epithet means: ‘from the west’; Linnaeus named tened, light orange-brown; wings 2, marginal, of
another species orientalis (‘from the east’, i.e. China), equal size and shape, surrounding seed but leaving
but this is now accommodated in a different genus. a notch at one or both ends, 1–1.5 mm wide, thin
membraneous, translucent.
991
Vernacular names
Taxonomic notes
Northern White-cedar, White-cedar, Eastern White-
cedar, Arbor-vitae, Swamp Cedar, Thuier cèdre, This species is morphologically similar throughout
Cèdre blanc its range. Frequent vegetative reproduction through
layering, especially in boggy environment, produces
clonal trees characteristically with a curved basal
Description
part of the trunk and standing in (semi-)circles.
Trees to 20–25(–30) m tall; trunk monopodial, erect,
often curved at base, to 1–1.5(–1.8) m d.b.h. Bark
Distribution
on trunks relatively thin, fissured, often twisted in
a long spiral around the trunk, exfoliating in very NE North America: from Lake Winnipeg and the
long, thin strips, grey-brown. Branches spreading; Great Lakes to the Canadian Maritime Provinces
lower branches curved down, then upward, per- and the Appalachian Mountains.
sistent; foliage branches drooping or subpendu- TDWG codes: 71 MAN 72 NBR NSC ONT PEI QUE 74
lous, forming a conical to pyramidal crown. Foliage ILL MIN WIS 75 CNT INI MAI MAS MIC NWH NWY
branches plagiotropic; higher order branchlets flat- OHI PEN RHO VER WVA 78 KTY NCA TEN VRG
tened, covered with leaves, 5–20 × 2–3 mm, green,
turning orange-brown with dying leaves, persistent.
Ecology
Leaves decussate, imbricate, appressed or laterals
with free apices, dimorphic, all scale-like, on (sub) This species occupies a geographical range touching
ultimate branchlets 1–4 × 1–2 mm; facials rhombic in the north on the subarctic taiga-tundra interface
to obtrullate; apex obtuse, acute or short mucro- and in the south it extends well into the belt of de-
nate; laterals slightly larger, partially covering base ciduous angiosperm forests. It is therefore associ-
of facials but spreading from their apex, bilaterally ated both with conifers, most common of which are
flattened, incurved; margins entire; apex connate, Abies balsamea, Larix laricina in the boreal zone,
acute or obtuse; primary stomatal bands abaxial, Picea mariana, L. laricina and Pinus banksiana in
on underside of laterals and on adjacent facials; swampy sites, and Pinus strobus and Tsuga canaden­
glands conspicuous on facial leaves on both sides of sis in uplands, and with angiosperms, mainly Popu­
branchlets near leaf apex; leaf colour green or grey- lus balsamifera, P. tremuloides, Betula papyrifera,
green, stomatal bands more or less whitish green, B. alleghaniensis, Acer rubrum, and Fraxinus nigra
glands yellowish. Pollen cones globose, 1–1.5 mm in the tree layer and Alnus rugosa, Acer spicatum,
diam., yellowish maturing brown; microsporophylls Cornus stolonifera, and Vaccinium spp. in the shrub
4–6, decussate, peltate, bearing 3–4 abaxial, relative- layer. The altitudinal range is (10–)150–600(–670)
ly large, yellow pollen sacs. Seed cones numerous, m a.s.l. It grows equally well in swamps and on dry
close together above pollen cones, terminal on up- ground, but avoids extremes of both habitats; it is
curved, quadrangular branchlets with rhombic, gib- often growing abundantly on soils over limestone
bous scale leaves, narrowly ovoid when closed, 8–12 in upland areas and on alluvial soils with a high
× 4–6 mm, erect at maturity. Bract-scale complexes organic and mineral content in lowlands (rich fens
(6–)8(–10), decussate; lowest pair much reduced; supporting forest). This species also invades aban-
higher pairs narrowly oblong; upper pair connate doned fields and pastures. The climate is cool to cold
 temperate and relatively moist, with a short growing readily layering, to 3–4(–6) m d.b.h. Bark on trunks
season especially in the north of its range. relatively thin, fissured, fibrous, exfoliating in long,
shredding strips, dark reddish brown, exposed bark
grey-brown. Branches spreading, with lower branches
Conservation
curved down, then upward; foliage branches droop-
IUCN: LC ing or subpendulous, forming a conical to pyramidal
crown. Foliage branches plagio­tropic; higher order
branchlets flattened, covered with leaves, 5–20 × 2–3
Uses
mm, green, turning reddish brown with dying leaves,
The rot resistant properties of the wood of this spe- persistent. Leaves decussate, imbricate, appressed or
992 cies, known as ‘northern white cedar’ by lumber- laterals with free apices, dimorphic, all scale-like,
men, make it an excellent timber for outdoor uses. on (sub-)ultimate branchlets 1–4 × 1–2 mm; facials
These include rustic fencing, garden gates and sheds, rhombic to obtrullate, weakly ridged; apex obtuse,
and shingles. Canoes are still made from this wood acute or short mucronate; laterals equally long or
because of its light weight, as are tubs and barrels. slightly larger, partially covering base of facials but
More industrial uses are found in kraft pulp and spreading from their apex, bilaterally flattened, in-
particleboard. The leaves were traditionally used in curved; margins entire; apex connate or free, acute;
Native American medicine to prepare a decoction to primary stomatal bands abaxial, on underside of lat-
suppress fevers and coughs and to cure rheumatism. erals and on adjacent facials; glands inconspicuous
White-cedar was introduced to Britain in 1596 and or absent on leaves of ultimate branchlets; colour of
has been widely planted as a decorative tree. In hor- upper leaves lustrous green or dark green, of lower
ticulture there are many cultivars in common use, leaves and bases of leaves glaucous green, stomatal
both tree forms and dwarfed forms. These empha- bands whitish green. Pollen cones ovoid-oblong,
size shape of crown (while young plants) and colour (2–)3–4 × 1.5–2 mm, purplish, maturing to black-
of foliage, the latter with many shades of yellow in ish grey; microsporophylls 8–14, decussate, peltate,
the new foliage. Some cultivars retain juvenile leaf bearing 3–4 abaxial, reddish yellow pollen sacs. Seed
shapes for many years looking like needles instead cones numerous, close together above pollen cones,
of the normal decussately arranged scales. This spe- terminal on short, upcurved, quadrangular branch-
cies can be used in hedges as it takes clipping well. lets with rhombic, gibbous scale leaves, narrowly
ovoid when closed, erect, 10–16(–18) × 6–8 mm.
Bract-scale complexes 8–12(–14), decussate; lowest
Thuja plicata Donn ex D. Don, in Lambert, Descr. pair much reduced; upper pairs small, irregularly
Pinus 2: [19] 1824. Type: Canada: British Columbia, linear; 2–3 largest pairs spreading, (6–)7–8 × 3–5
Vancouver Island, Nootka Sound, [“Longinquiter mm, (ob)ovate-oblong, thin, obtuse or acute, with
on the N. W. C. Am.”], A. Menzies s.n. [ex herb. a sub­apical, straight or recurved, 1 mm long acute
W. J. Hooker] (lectotype K). Fig. 343 bract tip, abaxially dull brown, adaxially yellowish
brown or reddish brown, lustrous. Seeds 4–10 per
cone, usually only 4–6 (well) developed, 4–5 × 1.5–2
Etymology
mm, flattened, light yellowish brown; wings 2, mar-
The species epithet plicata means ‘folded’ and refers ginal, surrounding seed but leaving a notch at one
to the shape of the lateral leaves. or both ends, 1–1.5 mm wide, thin membraneous,
translucent.
Vernacular names
Distribution
Western Red-cedar, Pacific Red-cedar, Giant Cedar,
Giant Arbor-vitae, Canoe Cedar, Shinglewood W North America: along the Pacific Coast Range
and Cascade Range from S Alaska to N California
and in the N Rocky Mountains from British Colum-
Description
bia to Idaho and W Montana.
Trees to 65–70(–75) m tall; trunk monopodial, erect, TDWG codes: 70 ASK 71 ABT BRC 73 IDA MNT WAS
often fluted or buttressed at base, or multistemmed, 76 CAL
 a decrease of occupancy of Thuja plicata. Plantation
Ecology
forestry focusing on this species should eventually
The two more or less disjunct areas in which this reduce the level of exploitation of natural stands, in
species occurs: Pacific coastal mountains and Rocky particular in ‘old growth’ forest with its high ecologi-
Mountains, experience a different climate and there- cal value. At present this species is not considered to
fore sustain different forest types. The mostly much be in danger of extinction.
wetter (winter rainfall, up to 6600 mm p.a.) and IUCN: LC
milder coastal ranges support the tallest conifer for-
ests in the world, with Sequoia sempervirens in the
Uses
southern part exceeding 110 m and with Abies gran­
dis to 80 m, A. procera 85 m, Picea sitchensis 87 m, The wood of this species provided the main build- 993
Pinus lambertiana 75 m, Pseudotsuga menziesii 100 ing material for the Amerindian tribes along the
m, and Tsuga heterophylla to 80 m tall. Many of these Pacific coast, who developed a technique to split
trees also exceed any of their congeners elsewhere in large planks from the lower boles of big trees with-
overall size (Van Pelt, 2001). Thuja plicata, with max. out destroying the trees themselves. Nowadays, its
75 m, is one of the longest-lived in these forests, with main use is for making shingles used in roofing
veteran trees often in excess of 1000 years. Other residential buildings; as in most Cupressaceae, the
conifers in these coastal forests are Chamaecyparis wood is decay-­resistant and easy to work. For large
lawsoniana (extreme southern part of range), Xan­ construction purposes it is less suitable as it tends to
thocyparis nootkatensis, Calocedrus decurrens, Abies split, but it can be used for a variety of smaller utili-
amabilis, Pinus monticola, Tsuga mertensiana, and ties from garden sheds, glass houses, and furniture
Taxus brevifolia in the understorey. Common an- to tools. Western Red-cedar has been used in for-
giosperm trees are Acer macrophyllum, Alnus rubra estry plantations in some countries in NW Europe
along rivers, and Populus trichocarpa; in the shrub on a rather limited scale; it requires high rainfall
layer are especially abundant Vaccinium spp., Ru­ and performs best in the wetter parts of the British
bus spectabilis and Ribes bracteosum. Deep layers Isles. Thuja plicata has been widely planted as an or-
of mosses and liverworts cover the forest floor and namental tree in parks and large gardens. It is also
lower sections of tree trunks as well as fallen logs, on suitable for hedges as it grows back quickly from
which latter most conifers find the only substrate to clipping. Fewer cultivars are known from this spe-
germinate. In the interior Abies grandis, A. lasiocar­ cies than from T. occidentalis, but it is nevertheless
pa, Larix occidentalis, Picea engelmannii, P. glauca, of substantial importance in the horticultural trade.
Pinus contorta, P. ponderosa, Pseudotsuga menziesii
var. glauca, and Taxus brevifolia are the most com-
monly associated conifers. Here annual precipita- Thuja standishii (Gordon) Carrière, Traité Gén.
tion does not exceed 1200 mm and winters are much Conif., ed. 2, 1: 108. 1867. Thujopsis standishii
colder than along the coast. The altitudinal range of Gordon, Pinetum Suppl.: 100. 1862. Type: Japan:
this species is 1–2100(–2300) m a.s.l. It grows on a Honshu, Kanagawa Pref., Yokohama, leg. ign. s.n.
wide range of soil types over nearly all available geo- (holotype K).
logical formations.
Etymology
Conservation
This species was named after John Standish, a nurs-
This species is common especially in the coastal sec- eryman at whose tree nursery it was first grown in
tions of its extensive range and somewhat less so in England.
the interior parts. (Selective) logging of mature trees
and ‘old growth’ forest in which this species is a co-
Vernacular names
dominant continues in many areas where the forest
is not on protected land. In situations where second- Japanese Arbor-vitae; Kurobe, Nezu, Nezuko (Japa-
ary forest growth is managed to favour other spe- nese)
cies (e.g. Pseudotsuga menziesii), this would lead to
 brata, Tsuga diversifolia, Picea jezoensis, Pinus parvi­
Description
flora, and Larix kaempferi. Common angiosperm
Trees to 20–25(–30) m tall; trunk monopodial, erect, trees are Betula ermanii, B. corylifolia, Fagus sie­
or multistemmed, to 1 m d.b.h. Bark on trunks rela- boldii, Quercus mongolica var. grosseserrata, and
tively thin, exfoliating in short, thin, curling flakes, Zelkova serrata. Thuja standishii is a minor compo-
reddish brown with grey-brown flakes. Branches nent in these forests, usually remaining a smaller
spreading or ascending, foliage branches drooping tree than the dominants and not exceeding 15 m in
or subpendulous, forming a pyramidal or broadly height. Often it is confined to moist rocky precipices
spreading crown. Foliage branches plagiotropic, facing north, sites less suitable for larger trees. Its
higher order branchlets flattened, covered with altitudinal range is (250–)500–2000(–2500) m a.s.l.
994 leaves, forming more or less rounded sprays, green,
abruptly turning reddish brown with dying leaves,
Conservation
persistent. Leaves decussate, imbricate, dimorphic,
all scale-like, on (sub)ultimate branchlets 1.5–5 × IUCN: LC
1–2.5 mm; facials rhombic to obtrullate; apex obtuse
or acute to weakly apiculate; laterals slightly larger,
Uses
partially covering base of facials but spreading from
their apex, bilaterally flattened, incurved; margins The wood of this species is highly prized for special
entire; apex connate, acute-apiculate; primary sto- building purposes, e.g. ceilings and panelling, and is
matal bands abaxial, on underside of laterals and on also used for furniture, fanlights and clogs (wooden
adjacent facials; glands inconspicuous on (larger) shoes). It was in the past regarded as one of the ‘five
facial leaves, small; leaf colour (yellowish) green to trees of Kiso’ or ‘Tome-ki’ (preserved trees) which
dark green, stomatal bands faintly whitish green. meant they were the property of the Emperors of
Pollen cones ovoid-oblong, 3–4(–5) mm long, pur- the Tokugawa dynasty in the 17th and 18th centuries
ple maturing purplish black; microsporophylls 8–16, and not to be used by common people. This prac-
decussate, peltate, bearing 3–4 abaxial, yellow pol- tice preserved several forests from overexploitation.
len sacs. Seed cones above pollen cones, terminal on In Japan it has been used as an ornamental tree for
short, straight, quadrangular branchlets with rhom- many centuries. Robert Fortune first saw it in 1860
bic, gibbous scale leaves, broadly ovoid, 7–12(–14) in gardens in Tokyo and introduced it to England
× 6–7 mm, turning yellowish brown. Bract-scale via Standish’s nursery at Ascot. It is still in cultiva-
complexes 8–10(–12), decussate; lowest pair much tion in Europe and the USA but much less common
reduced; higher pairs irregular, partly spreading; there than some of the other species and mostly seen
two largest pairs ca. 5–7 × 4–5 mm, ovate-rhombic as specimen trees in dendrological collections.
to obtrullate, thin, obtuse or acute, with a subapical
triangular bract tip of ca. 0.5 mm. Seeds (6–)8–10
per cone, 4–5 × 2 mm, flattened, tapering towards Thuja sutchuenensis Franch., J. Bot. (Morot) 13 (8):
base, light brown; wings 2, marginal, surrounding 262. 1899. Type: China: Chongqing Municipality,
seed but leaving a notch at micropylar end, 1 mm Chengkou Xian, Daba Shan, [“Tchen-keou-tin”
wide, thin membraneous, apically puberulent, light (vicinity of Chengkou)], P. G. Farges 1158 (holotype
brown. P). Fig. 344

Distribution Etymology
Japan: Honshu, Shikoku. The species epithet derives from Sichuan, China, in
TDWG codes: 38 JAP-HN JAP-SH which the locality where the species was found at
the time was located; this area has now been sepa-
rated from that province.
Ecology
Thuja standishii occurs in mixed montane conifer
Vernacular names
or conifer-angiosperm forests, with Abies homolepis,
Chamaecyparis obtusa, C. pisifera, Thujopsis dola­ ya bai shu
 TDWG codes: 36 CHC-CQ
Description
Trees to 20 m tall (sometimes a shrub); trunk mo-
Ecology
nopodial, terete, to 30 cm d.b.h. Bark on trunks to
4–5 cm thick, fibrous and fissured lower down, ex- This very restricted species occupies steep slopes
foliating in short, thin, curling flakes, cinnamon-red and ridges of limestone mountain sides between
or reddish brown to grey-brown. Branches spread- 800 m and 2100 m a.s.l. in mixed angiosperm shrub-
ing or ascending, forming a conical or pyramidal land and forest. The soil is mountain yellow-brown
crown. Foliage branches plagiotropic; higher order earth developed from limestone, rich in organic
branchlets flattened, covered with leaves, pinnately matter (2.3–3%) and with a thick humus layer (ca.
arranged up to ultimate branchlets, forming fron- 20 cm) and a pH of 6.2–7. The climate is warm tem- 995
dose sprays, green, abruptly turning orange-brown perate and humid with mean annual precipitation
with dying leaves, persistent. Leaves decussate, ca. 1400 mm.
imbricate, dimorphic, all scale-like, on (sub)ulti-
mate branchlets 1.5–4 × 1–1.5 mm; facials rhombic
Conservation
to obtrullate, weakly keeled; apex obtuse; laterals
slightly larger, broadly falcate, partially covering Formerly this taxon was listed as the only conifer
base of facials but spreading from their apex, bilater- Extinct in the Wild (EW) by IUCN-SSC (Farjon &
ally flattened, incurved; margins entire; apex obtuse; Page, 1999). In October 1999 it was rediscovered by a
primary stomatal bands abaxial, on the underside regional team of Chinese botanists, not far from (or
of laterals and on the adjacent facials; leaves ap- perhaps even at) the locality where R. P. Farges had
parently eglandular, lustrous green, stomatal bands made his last collections almost 100 years previously
whitish green or glaucous. Pollen cones subglobose, (Xiang et al., 2002). There is evidence of cutting at
2–3 mm long, yellowish green maturing brown; mi- the locality, and presumably the remaining individ-
crosporophylls 6–8, decussate, peltate, bearing (2–)3 uals are only those left in inaccessible places and/or
abaxial, yellow pollen sacs. Seed cones terminal on of small stature. Seedlings are very rare so there is
short, straight branchlets with rhombic, gibbous serious concern about regeneration capacity of the
scale leaves, (ob)ovoid or elliptical, 5–8 × 3–4.2 mm, remnant population. In order to achieve better pro-
with slightly spreading scales. Bract-scale complexes tection, work involving local stakeholders is being
8–10, decussate; lowest pair reduced; highest pair(s) carried out with help from the Bedgebury Pinetum
small, connate; two largest pairs ca. 3–5 mm long, (UK Forestry Commission) in Kent, England.
obovate, thick, obtuse, with a subapical ridge and IUCN: CR (B1 + 2b,c,e)
triangular bract tip 0.5–1 mm, dull brown abaxially,
lustrous yellowish brown or orange-brown adaxi-
Uses
ally. Seeds ca. 4 per cone, 3–4 × 1.5 mm, flattened,
tapering towards both ends, lustrous light brown; The wood of this species is soft, light, easily worked
wings 2, marginal, surrounding seed but leaving a and durable. It is used for applications requiring de-
notch at both ends, 0.5 mm wide, thin membrane- cay resistance by local people, e.g. home construc-
ous, yellowish brown. tion, production of shingles, application for funeral
services etc. It is too rare to possess much commer-
cial value and outside China this species is not in
Distribution
cultivation.
China: Chongqing (Chengkou Co., Kaixian Co.), Si-
chuan (Wangyuan Co.), on the southern slope of the
Dabashan Mountains.
 Thujopsis Siebold & Zucc. ex Endl., Gen. Pl. Suppl. 2: 24. 1842 (nom. cons.).
Type: Thujopsis dolabrata (Thunb. ex L. f.) Siebold & Zucc. [Thuja dolabrata L. f.]
(Cupresaceae).

Dolophyllum Salisb., Quart. J. Sci. Lit. Arts 2 (4): 313. margins entire; apex free, incurved, obtuse or acute;
1817 (nom. rej.). Type: Dolophyllum dolabrata (L. f.) the broad primary stomatal bands all on underside
Salisb. [Thuja dolabrata L. f.]. of plagiotropic shoots, on both sides on erect shoots;
leaf colour above lustrous dark green; stomatal
Greek: opsis = ‘aspect’, ‘appearance’, i.e. appearing bands conspicuous, white, bordered by light green
996 similar to the genus Thuja. margins. Pollen cones terminal, solitary, ovoid-
globose, 3–4 mm long, reddish purple maturing to
red-brown; microsporophylls 8–10, decussate, pel-
Description
tate, rounded, bearing 3–5 abaxial pollen sacs. Seed
See the species description. cones terminal on straight branchlets with small but
unmodified scale leaves, subglobose, markedly um-
bonate, 8–16(–20) mm long. Bract-scale complexes
Distribution
6–8(–10), decussate; lower and upper pairs reduced;
As for the species. 4 larger pairs cuneate or obdeltoid, apically thick-
ened, subapically reflexed with a large 2–3 mm long,
emerging and upturned bract tip, abaxially rugose,
Thujopsis dolabrata (Thunb. ex L. f.) Siebold & dull brown to grey, adaxially striated, with 2–5 seed
Zucc., Fl. Japon. 2 (4): 34, tt. 119, 120. 1844. scars at base, light brown. Seeds 15–25 per cone, nar-
rowly ovate, irregularly compressed, 4–5 × 2 mm,
greyish brown; wings 2, lateral, 1–1.5 mm wide, thin
Etymology
menbranous, yellowish.
The species epithet, from Latin dolabra = hatchet or
mattock, refers to the shape of the facial leaves.
Distribution
Japan: Hokkaido, Honshu, Kyushu, Shikoku.
Vernacular names
TDWG codes: 38 JAP-HK JAP-HN JAP-KY JAP-SH
Hiba Arbor-vitae; Asurano, Asuhi, Hiba (Japanese)
Ecology
Description
This species is a constituent of conifer and conifer-
Trees to 15–20(–30) m tall, sometimes a large shrub, angiosperm forests from lowland coastal areas to
evergreen, monoecious; trunk monopodial or mul- montane sites in regions with a cool, moist climate.
tistemmed, to 1 m d.b.h.; branches frequently layer- Its altitudinal range is 400–1800(–2100) m a.s.l. The
ing. Bark on trunks with longitudinal plates, reddish most commonly associated conifer is Tsuga diversi­
brown with greyish strips of outer bark. Branches folia, but it can grow in natural forests with sev-
spreading or ascending to erect in (layering) shrubs, eral others, e.g. Chamaecyparis obtusa, C. pisifera,
forming pyramidal to broadly conical or domed Crypto­meria japonica (locally), Thuja standishii,
crowns. Foliage branches in flat but irregular sprays, Sciado­pitys verticillata (locally), Abies homo­lepis,
branching alternately or divaricately; small branch- A. mariesii, and Pinus parviflora. Common an-
lets variable in length, distinctly flattened, persistent. giosperm trees are Betula ermanii, Fagus sieboldii,
Leaves decussate, imbricate, dimorphic (on older, Quercus mongolica var. grosseserrata, Pterocarya
thickening branches becoming monomorphic, de- rhoifolia, Aesculus turbinata, and Cercidiphyllum
current and gradually widening with the growing japonicum. Only occasionally does this species form
shoot), (1–)3–8 × 1–5 mm on flattened branchlets; more or less pure stands, this is mainly the case with
facials dolabriform or spathulate, obtuse; laterals the northern variety T. dolabrata var. hondae. Else-
spreading from below their apex, bifacially flattened; where, trees are mostly scattered and small and must
be extremely shade tolerant, growing under canopy mm long, bract-scale complexes distinctly um-
of Tsuga or Fagus and competing with e.g. Ilex rugo­ bonate.
sa (Wilson, 1916). The variable disposition of foliage
branches and the development of stomatal zones on
Distribution
leaves indicate adaptability to changing light condi-
tions; the capacity of layering is probably another Japan: S Honshu, Kyushu, Shikoku.
strategy to maintain sufficient light interception TDWG codes: 38 JAP-HN JAP-KY JAP-SH
under an expanding canopy, especially in earlier
phases of growth.
Conservation
IUCN: LC 997
Uses
Hiba Arbor-vitae is an important forest tree in Japan Thujopsis dolabrata (Thunb. ex L. f.) Siebold &
and is one of the ‘five trees of Kiso’ (all are conifers) Zucc. var. hondae Makino, Bot. Mag. (Tokyo) 15:
originally preserved for imperial use. There are now 104. 1901 [“hondai”]. Type: Japan: Honshu, Aomori
managed state forests with this species as the domi- Pref., Aomori, [“Prov. Mutsu, near Awomori”],
nant tree besides some imperial forests. Its light, soft, T. Makino s.n.(?) (holotype TI).
resinless and durable wood is used in construction,
for bridges, buildings, furniture, the wooden basis of
Description
laquer work, wood carving, etc. Younger trees of this
species are valued for their ornamental qualities due Foliage relatively slender, with on average slightly
to a striking contrast between upper lustrous green smaller leaves than in var. dolabrata. Seed cones 15–
and lower white banded leaf surfaces. Hiba Arbor- 20 mm long, bract-scale complexes less umbonate.
vitae is widely planted in Japan as well as in other
countries with a temperate maritime climate. Sever-
Distribution
al cultivars are known, most common are those with
variegated leaves and some dwarf forms with com- Japan: Hokkaido, N Honshu.
pact growth and smaller leaves. The earliest of these TDWG codes: 38 JAP-HK JAP-HN
were selected in Japan and introduced to Europe at
the same time as the species, around the middle of
Conservation
the 19th century.
IUCN: LC
2 varieties are recognized:

Thujopsis dolabrata (Thunb. ex L. f.) Siebold &

Zucc. var. dolabrata. Thuja dolabrata Thunb. ex
L. f., Suppl. Pl.: 420. 1782. Type: Japan: Honshu
[“Habitat in Japonia”], C. P. Thunberg UPS 22555
(holotype UPS). Fig. 345, 346

Description
Foliage relatively coarse with on average slightly
larger leaves than in var. hondae. Seed cones 8–15
 Torreya Arn., Ann. Mag. Nat. Hist., ser. 1, 1: 130. 1838 (nom. cons.). Type: Torreya
taxifolia Arn. (Taxaceae).

Caryotaxus Henkel & W. Hochst., Syn. Nadelhölz.: 1b. Leaves 1.2–3.8(–4.5) cm long. Branches spread-
366. 1865. Type: Caryotaxus nucifera (L.) Henkel & ing and ascending 3
W. Hochst. [Taxus nucifera L.]; Foetataxus J. Nel- 2a. Leaves mostly drooping; stomatal bands 0.5–1
son, Pinaceae: 167. 1866. Type: Foetataxus montana mm wide; margins on that side of leaves 0.5–0.7
J. Nelson (nom. illeg.) [Torreya taxifolia Arn.]; Tumi­ mm wide. Ripe fleshy aril including seed 10–12
on Raf. ex Greene, Pittonia 2: 193. 1891. Type: Tumion mm wide, pruinose to reddish yellow T. jackii
998 taxifolium (Arn.) Greene [Torreya taxifolia Arn.]. 2b. Leaves pectinately arranged; stomatal bands
0.3–0.4 mm wide; margins 0.7–1 mm wide.
Named after John Torrey (1796–1873), American Ripe fleshy aril including seed to 25 mm wide,
botanist. green with purplish streaks T. californica
3a. Leaves straight or slightly down-curved, with a
long cuspidate or spinose apex T. nucifera
Description
3b. Leaves straight or distally (slightly) falcate, with
Dioecious or rarely monoecious evergreen trees a short cuspidate apex 4
or sometimes shrubs. Resin canals in leaves (1) 4a. Leaves (0.7–)1–2.7(rarely to 4.5) cm long; sto-
and seed arils. Bark smooth or flaking, becom- matal bands 0.3–0.8 mm wide. Ripe fleshy aril
ing fissured. Leaves alternate (helically inserted), including seed pale purplish brown or whitish
pectinately arranged, flattened, linear-lanceolate, pruinose T. grandis
rigidly coriaceous, asperous, sharply acuminate or 4b. Leaves 1.2–3.8(–4.5) cm long; stomatal bands
pungent, with a prominent midrib. Stomata in two 0.3–0.5 mm wide. Ripe fleshy aril including
separated bands on abaxial side. Pollen cones soli- seed green streaked with purple, or slightly
tary in the axil of each leaf on fertile shoots, form- whitish pruinose 5
ing double rows on the underside of plagiotropic 5a. Midrib on the abaxial (lower) side of leaves
foliage branchlets, subglobose; microsporophylls 0.3–0.6 mm wide, leaf margins on that side
decussately arranged, tightly clustered, peltate, with 0.5–1.2 mm wide, total leaf width 2–4 mm. Ripe
3–6 pendulous pollen sacs forming a small umbel fleshy aril including seed pale green or slightly
and containing subspherical, arillate pollen. Seed-­ whitish pruinose T. fargesii
bearing structures axillary to foliage leaves, consist- 5b. Midrib on the abaxial side of leaves 0.6–0.8
ing of a miniaturized branching shoot with a few mm wide, leaf margins on that side 0.6–0.8 mm
spirally inserted bracts on the main and lateral axes wide, total leaf width 2–3.2 mm. Ripe fleshy aril
and usually 2–3 sub-terminal, erect ovules of which including seed dark green streaked with pur-
only 1 develops into a large, hard seed with a strong- ple T. taxifolia
ly sclerified seed coat and completely surrounded
(except at the distal tip) by a fleshy aril which ripens
in the second year and becomes purplish green to Torreya californica Torr., New York J. Pharm. 3: 49.
bluish black. Seedlings with 4 short cotyledons. 1852. Type: not designated. Fig. 347, 348
6 species.
Etymology
Distribution
The species epithet alludes to its Californian native
Asia: China, S Korea, Japan. North America: USA range.
(disjunct): California, Florida.
Vernacular names
Key to the species of Torreya
California nutmeg, Stinking cedar
1a. Leaves (2–)3–6(–9) cm long. Branches spread-
ing and drooping 2
 to mountainous terrain dominated by tall conifers,
Description
especially Sequoia sempervirens, Picea sitchensis and
Trees (sometimes shrubs) to 25(–40) m tall; trunk Pseudotsuga menziesii. In the western valleys of the
to 1.2 m d.b.h. Bark irregularly vertically fissured, Sierra Nevada it is associated with angiosperms,
sometimes flaking, pale reddish brown or greyish mainly Acer spp., Cornus nuttallii, Platanus occiden­
brown, weathering to dark grey. Branches spreading talis, and Alnus rubra and here it strictly adheres to
or somewhat drooping, forming a broad conical to stream banks in steep canyons at altitudes around
rounded crown. Foliage branchlets slender, usually 1200 m a.s.l. Its altitudinal range is from near sea
with opposite branching, sometimes with more than level (but usually above 200 m) in the Coast Ranges
2 laterals from a node, spreading horizontally at 40– to 2500 m a.s.l. in the Sierra Nevada. On serpentine
60° from the main axis in most cases, terete, with rock it becomes a stunted tree or shrub and occurs 999
grooves along margins of decurrent leaf bases, green on N-facing slopes in coastal chaparral.
turning reddish brown after the first year. Buds at
ends of branchlets very small; bud scales at lower
Conservation
node of previous year enlarged, triangular, keeled,
lustrous brown, deciduous thereafter. Leaves pec- IUCN: NT
tinately arranged, spreading at 45–90° from shoot
axis, linear to linear-lanceolate, (2–)3–6(–8) cm long,
Uses
straight or slightly falcate, 2.4–4.5 mm wide, with
1–2 mm long, twisted petiole, abruptly widening at Torreya californica is the most commonly cultivated
base, tapering to a long cuspidate apex, coriaceous, species of its genus and can grow into an impressive,
lustrous green on adaxial (upper) side, smooth, flat. wide crowned tree with striking foliage and large,
Stomatal bands on abaxial side only, very narrow pendulous, plum-like green or glaucous seeds. It is
(0.3–0.4 mm), greenish white or glaucous, separated almost exclusively used as an ornamental in large
by a raised, 0.6–1.2 mm wide, green midrib and bor- gardens or planted in arboreta. In the past, its wood
dered by 0.7–1 mm wide, raised, flat margins; stoma- was used for fencing and bridges, as it is rot resist-
ta small, randomly distributed. Pollen cones axillary, ant. California Indians used it for their bows. The
solitary, forming short rows on underside of lower name ‘nutmeg’ only refers to a superficial similarity
leaves of lateral branchlets, with several pairs of ba- with true nutmeg fruits (from Myristica fragrans, an
sal bud scales in 4 rows, ca. 5–6 × 4.5–5 mm before angiosperm tree species). The native tribes roasted
anthesis, pale yellow or nearly white; microsporo­ the seeds for food and some of them (e.g. the Pomo
phylls numerous, peltate, each with 4–6 marginal, tribe) used the roots for basket weaving.
pendulous pollen sacs. Seed bearing structures
axillary, solitary or in pairs, sessile, with rounded,
keeled bract scales subtending seed. Ripe fleshy Torreya fargesii Franch., J. Bot. (Morot) 13 (8): 264.
aril including seed smooth, light green or glaucous 1899.
green, with darker green to purplish longitudinal
streaks from base to below apex, obovoid or broadly
Etymology
elliptic, 25–35 mm long, to 25 mm wide, mucronate
at apex. Seed proper smooth or with 2 opposite lon- This species has been named after the French mis-
gitudinal ridges. sionary and plant collector P. G. Farges.

Distribution Vernacular names

SW USA: California (Coast Ranges, Sierra Nevada). Farges nutmeg tree; ba shan fei shu (Chinese)
TDWG codes: 76 CAL
Description
Ecology
Shrubs or trees to 20 m tall; trunk to 1 m d.b.h. Bark
Torreya californica is a scattered understorey tree irregularly vertically fissured, sometimes flaking,
usually growing in moist and shady places in hilly pale brown or greyish brown, weathering to dark
 grey. Branches spreading and ascending, forming than the margins; in T. grandis midrib and margins
a broad crown. Foliage branchlets slender, usu- are of about equal width and as wide as the margins
ally with opposite branching, sometimes with more in T. fargesii. With roughly equal total leaf width in
than 2 laterals from a node, spreading horizontally both species, this results in wider stomatal bands in
at 40–60° from main axis in most cases, terete, with T. fargesii. Two varieties are recognized also with this
grooves along margins of decurrent leaf bases, green species: var. fargesii and var. yunnanensis; the latter
turning yellowish brown after the first year. Buds at has longer and wider leaves than both T. fargesii var.
ends of branchlets very small; bud scales at lower fargesii and T. grandis, but retains the relative widths
node of previous year enlarged, broadly triangular, mentioned above on the abaxial leaf faces. This char-
keeled, lustrous brown, deciduous thereafter. Leaves acter state is therefore accepted as sufficient to sepa-
1000 pectinately arranged, spreading at 55–90° from rate T. fargesii and T. grandis at species level.
shoot axis, linear to linear-lanceolate, 1.2–3.5(–4.5)
cm long, straight or distally falcate, 2–4 mm wide,
Distribution
with 0.5–1 mm long, twisted petiole, abruptly wid-
ening at base, tapering to a cuspidate apex, coria- China: Chongqing, S Anhui (?), W Hubei, NW Hu-
ceous, lustrous green on adaxial (upper) side, with nan, Jiangxi, S Shaanxi, Sichuan (Emei Shan [Mt.
two grooves on either side of an indistict midvein Omei]), NW Yunnan.
at least in lower half of blade. Stomatal bands on TDWG codes: 36 CHC-CQ CHC-HU CHC-SC CHC-
abaxial side only, 0.3–0.5 mm wide, light brown, YN CHN-SA CHS-HN
separated by a 0.3–0.6 mm wide, green midrib and
bordered by 0.5–1.2 mm wide, flat or very narrowly
Ecology
revolute green margins; stomata small, randomly
distributed. Pollen cones axillary, solitary, forming This species occurs in coniferous, mixed and broad-
short rows on underside of lower leaves of lateral leaved forests as an understorey shrub or a small to
branchlets, with several pairs of basal bud scales in medium sized tree, in mountains from 1000 m to
4 rows, ca. 5–6 × 4.5–5 mm before anthesis, pale yel- 3400 m a.s.l.
low; microsporophylls numerous, peltate, each with
(3–)4–5 marginal, pendulous pollen sacs. Seed bear-
Uses
ing structures axillary, in pairs, sometimes grouped
together on a branchlet, sessile, with rounded, keeled This species yields high quality wood used in build-
bract scales subtending seed. Ripe fleshy aril includ- ing houses, bridges (durability of the wood) and fur-
ing seed smooth, pale green or slightly whitish pru- niture; some of it is also used to make utensils and for
inose, ovoid, globose or broadly ellipsoid, 15–25 mm wood turning. The seeds are edible and produce oil
diam., mucronate at apex. Seed proper smooth or that is extracted; the essential ‘torreya oil’ is extract-
with 2 opposite longitudinal ridges. ed from the aril. It is in cultivation in China, but not
as widely as T. grandis; outside its native country it is
only planted in a few botanic gardens and arboreta.
Taxonomic notes
This species, treated as a variety of Torreya grandis 2 varieties are recognized:
in the World Checklist and Bibliography of Conifers
(Farjon, 1998, [2001]) and by most western botanists, Torreya fargesii Franch. var. fargesii. Torreya
is recognized as a distinct species in Flora of China grandis Fortune ex Lindl. var. fargesii (Franch.)
4: 95 (1999). In that Flora, T. grandis has two varie- Silba, Phytologia Mem. 7: 74. 1984. Types: China:
ties, one (var. grandis) widespread in eastern China Chongqing Municipality, Chengkou Xian, Daba
and one (var. jiulongshanensis) limited to S Zhe- Shan [near “Heou pin”], P. G. Farges 100 & 945
jiang. Torreya fargesii partly overlaps that range, but (syntypes P).
extends further west and north. A consistent mor-
phological difference is found in the relative width of
Description
stomatal bands and green margins and midrib on the
leaf under (abaxial) sides, which appears to separate Leaves straight or rarely distally falcate, 1.2–2.5 cm
the two species. In T. fargesii the midrib is narrower long, 2–3.5 mm wide; grooves on adaxial surface
prominent only below middle of blade, often fading ­ opulations in Gongshan and Weixi are still of con-
p
towards shortly tapered apex. siderable size and should be conserved inside pro-
tected forest reserves.
IUCN: DD
Distribution
China: S Anhui (?), W Hubei, NW Hunan, Jiangxi,
S Shaanxi, Sichuan. Torreya grandis Fortune ex Lindl., Gard. Chron.
TDWG codes: 36 CHC-CQ CHC-HU CHC-SC CHN- 1857: 788. 1857.
SA CHS-HN
Etymology
1001
Conservation
The species epithet means ‘great’ and may refer to
This taxon is considered vulnerable on account of the potentially large size of the arillous seed, espe-
deforestation in many areas where it occurs (or has cially in some cultivated forms.
occurred in the recent past according to herbarium
collections made since the beginning of the 20th cen-
Vernacular names
tury).
IUCN: VU (A1c) Chinese nutmeg tree; fei shu, yuan bian zhong (Chi-
nese)
Torreya fargesii Franch. var. yunnanensis (W. C.
Cheng & L. K. Fu) N. Kang, Bull. Bot. Res. Harbin
Description
15 (3): 353. 1995. Torreya yunnanensis W. C. Cheng
& L. K. Fu, Acta Phytotax. Sin. 13 (4): 87. 1975; Tor­ Trees to 25 m tall; trunk to 1(–2) m d.b.h. Bark irreg-
reya grandis Fortune ex Lindl. var. yunnanensis ularly vertically fissured, sometimes flaking, yellow-
(W. C. Cheng & L. K. Fu) Silba, Phytologia 68: 72. ish grey or greyish brown, weathering to dark grey.
1990. Type: China: NW Yunnan [cited in Chinese Branches spreading and ascending, forming a broad
characters] Chinese collector 20946 (holotype PE). crown. Foliage branchlets slender, usually with op-
posite branching, sometimes with more than 2 lat-
erals from a node, spreading horizontally at 40–70°
Description
from main axis in most cases, terete, with grooves
Leaves straight or often distally falcate, (1.5–)2–4 cm along margins of decurrent leaf bases, green turn-
long, 3–4.5 mm wide; grooves on adaxial surface ex- ing yellowish green and then grey after first year.
tending to (near) apex. Buds at ends of branchlets very small; bud scales at
lower node of previous year enlarged, broadly tri-
angular, keeled, lustrous brown, deciduous there-
Distribution
after. Leaves pectinately arranged, spreading at
China: NW Yunnan (Gongshan, Lijian, Weixi, 65–90° from shoot axis, linear to linear-lanceolate,
Zhongdian). (0.7–)1–2.7(–4.5) cm long, straight or distally fal-
TDWG codes: 36 CHC-YN cate, 2–3.5(–4) mm wide, with a twisted, 0.5–1 mm
long petiole, abruptly widening at base, tapering to
a cuspidate apex, coriaceous, lustrous green on the
Conservation
adaxial (upper) side, with (or without) an indistict
This taxon was not earlier evaluated under IUCN midvein at least in lower half of blade. Stomatal
criteria; it can be assumed to be in a threatened bands on abaxial side only, 0.3–0.8 mm wide, light
category since its distribution is much more lim- brown, separated by a 0.4–0.7 mm wide, green mid-
ited than that of var. fargesii, while at least in some rib and bordered by 0.5–0.8 mm wide, flat or very
areas of NW Yunnan where it occurs deforestation narrowly revolute green margins; stomata small,
and forest disturbance may well have made a similar randomly distributed. Pollen cones axillary, solitary,
impact on the populations. It was considered vul- forming short rows on underside of lower leaves of
nerable in China Plant Red Data Book 1 (Fu & Jin, lateral branchlets, with several pairs of basal bud
1992) in which it was treated as a full species. The scales in 4 rows, ca. 5–8 × 4.5–5 mm before ­anthesis,
 pale yellow; ­microsporophylls numerous, peltate, streams and/or in shady spots in the forest, but large
each with 4–5 marginal, pendulous pollen sacs. trees can reach the canopy and compete well with
Seed bearing structures axillary, solitary or in pairs, many other tree species. In secondary vegetation it
sometimes grouped together on a branchlet, sessile, may persist for some time, but eventually disappears
with rounded, keeled bract scales subtending seed. due to excessive growth of bamboo and other weedy
Ripe fleshy aril including seed smooth, pale purplish plants.
brown or slightly whitish pruinose, obovoid-conical
to ovoid, or ellipsoid to oblong-ellipsoid, 20–40 mm
Uses
long, 12–25 mm wide, mucronate at apex. Seed prop-
er smooth or with 2 opposite longitudinal ridges. This species yields high quality wood used in build-
1002 ing houses, bridges (durability of the wood), and
furniture; some of it is also used to make utensils
Taxonomic notes
and for wood turning. The seeds are edible and pro-
Recently, a new species, T. parvifolia, was described duce oil that is extracted; the essential ‘torreya oil’ is
from Sichuan in China, the type specimens of extracted from the aril. The seeds proper (kernels)
which are deposited in local Chinese herbaria. Its are sold in Chinese drug shops as a remedy against
description, with an illustration, made comparison coughs and can be eaten as nuts. This species is com-
with T. yunnanensis, but not with T. grandis or any mon in cultivation in China (especially in Zhejiang),
other species. Its reported characters fit well within where some forms with various shapes and sizes (up
those given here and elsewhere (e.g. Flora of China to 5 cm long) of the arillous seeds are treated as cul-
4, 1999) for T. grandis var. grandis and it is there- tivars and these are planted as fruit trees or shrubs
fore treated here as a synonym of that taxon. A fair in special plantations. The Chinese nutmeg tree was
number of varieties and forms have been described introduced to England by Robert Fortune in 1855,
in this species; several of these are based on cultivat- but remains uncommon in cultivation in Europe
ed plants in China, which would probably be more and elsewhere. Where it is grown it develops into a
appropriately be termed cultivars. They are often shrubby habit, not a monopodial tree.
based on characters of the arrillous seeds and some
of these forms attain large sizes and are of economic 2 varieties are recognized:
importance in Chinese traditional medicine. Only
one recently described variety has been recognized Torreya grandis Fortune ex Lindl. var. grandis.
in Flora of China 4: 95 (1999); its seeds are differ- Torreya nucifera (L.) Siebold & Zucc. var. grandis
ently shaped and its leaves are longer than in other (Fortune ex Lindl.) Pilg., in Engler, Pflanzenr. IV.5
varieties. It is accepted here. [18]: 107. 1903. Type: China: Zhejiang, [“mountains
of Chekiang, China”], R. Fortune s.n. (holotype not
located, isotype K).
Distribution
China: Anhui, Chongqing, Fujian, Guizhou, Hubei, Torreya parvifolia T. P. Yi, L. Yang & T. L. Long, Bull.
Hunan, Jiangsu, Jiangxi, S Shaanxi, Sichuan, Zhe- Bot. Res. (China) 26 (5): 514. 2006.
jiang.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC-
Description
SC CHC-YN CHN-SA CHS-AH CHS-FJ CHS-HN CHS-
JS CHS-JX CHS-ZJ Leaves (0.7–)1–2.7 cm long. Ripe fleshy aril includ-
ing seed obovoid to ovoid or ellipsoid.
Ecology
Distribution
Torreya grandis is a constituent of the diverse mixed
mesophytic forest (remnants) in Zhejiang, in which China: Anhui, Fujian, Guizhou, Hunan, Jiangsu,
broad-leaved trees (angiosperms) dominate but Jiangxi, Zhejiang.
several conifers can be present as associated trees. TDWG codes: 36 CHC-GZ CHS-AH CHS-FJ CHS-HN
The altitudinal range of this species is between 200 CHS-JS CHS-JX CHS-ZJ
m and 1400 m a.s.l. It is commonly found along
 lower node of previous year enlarged, broadly trian-
Conservation
gular, keeled, lustrous brown, deciduous thereafter.
IUCN: LC Leaves mostly drooping, linear to linear-lanceolate,
(2.5–)3.5–7(–9) cm long, straight or slightly falcate,
Torreya grandis Fortune ex Lindl. var. jiulongshan­ (2.5–)3–4 mm wide, with a 1–2 mm long, twisted
ensis Z. Y. Li, Z. C. Tang & N. Kang, Bull. Bot. Res. petiole, abruptly widening at base, gradually taper-
Harbin 15 (3): 356. 1995. Type: China: Zhejiang, ing to a cuspidate apex, soft coriaceous, lustrous
Suichang, Jiulong Shan, Z. Y. Li & Z. C. Tang 9009 green on adaxial (upper) side, with an indistict mid-
(holotype PE). vein lying in a shallow depression at least in lower
half of blade. Stomatal bands on abaxial side only,
silvery grey to light brown, 0.5–1.4 mm wide, sepa- 1003
Description
rated by a 1–1.2 mm wide, centrally sharply raised
Leaves 2.5–4.5 cm long. Ripe fleshy aril including green midrib and bordered by 0.5–0.7 mm wide,
seed obovoid-conical. flat or slightly downcurved green margins; stomata
small, randomly distributed. Pollen cones axillary,
solitary, with several pairs of basal bud scales in
Distribution
4 rows, ca. 5–8 × 4.5–5 mm before anthesis, pale yel-
China: S Zhejiang (Suichang Xian). low; microsporophylls numerous, peltate, each with
TDWG codes: 36 CHS-ZJ 4–5 marginal, pendulous pollen sacs. Seed bearing
structures axillary, in pairs, sessile, with 2 pairs of
rounded, keeled bract scales and a small lateral bract
Conservation
subtending seed. Ripe aril including seed smooth,
IUCN: LC whitish pruinose or glaucous, ripening to reddish
yellow, obovoid, 20–30 mm long, 10–12 mm wide,
mucronate at apex. Seed proper smooth or rugose.
Torreya jackii Chun, J. Arnold Arbor. 6: 144. 1925.
Type: China: Zhejiang, Xianju, R. C. Ching 1779
Taxonomic notes
(lectotype PE).
This species has a morphological similarity to Ce­
phalotaxus fortunei when only sterile foliage is ob-
Etymology
served.
The species epithet commemorates the American
botanist William Jack (1795–1822).
Distribution
China, N Fujian (Taining, Puchen), NE Jiangxi, S
Vernacular names
Zhejiang.
Jack’s nutmeg tree; chang ye fei shu (Chinese) TDWG codes: 36 CHS-FJ chs-jx CHS-ZJ

Description Ecology
Shrubs or small trees to 12 m tall; trunk to 20 cm Torreya jackii occurs in evergreen broad-leaved forest,
d.b.h. Bark flaking, exposing pale brown new bark, along streams, on steep slopes in shade or in second-
weathering to grey or dark grey. Branches spread- ary vegetation usually near moisture. Its altitudinal
ing and drooping, forming a thin, open crown. range is between 400 m and 1000 m a.s.l. It is a species
Foliage branchlets slender, long and flexible, usu- of warm temperate to subtropical regions affected by
ally with opposite branching, sometimes with more the SE monsoon with an annual precipitation of 1350
than 2 laterals from a node, spreading or drooping at to 1600 mm and mean annual temperature 17–20 °C,
30–60° from the main axis in most cases, terete, with with absulute minimum of –10 °C. It grows on acidic
fine grooves along margins of decurrent leaf bases, mountain yellow earth or rocky granitic or rhyolitic
green turning lustrous reddish brown after first year. substrates with a low pH of 4.2–5 with good access
Buds at ends of branchlets very small; bud scales at to ground ­water or ­seepage. ­Associated species are
 e.g. Quercus oxy­phylla, Q. ­phillyrhaeoides, Photinia
Description
benthamiana, Loropetalum chinense, Cyclabalanop­
sis glauca, Castanopsis eyrei, Schima superba, and Shrubs or trees to 25 m tall; trunk to 1(–2) m d.b.h.
Rhododendron latoucheae. In secondary vegetation Bark irregularly vertically fissured, sometimes
shrubs like Loropetalum chinense, Vaccinium brac­ flaking in narrow strips, yellowish grey or greyish
teatum, Rhododendron ovatum, Symplocos caudata, brown, weathering to dark grey. Branches spread-
etc. are common. Roots have a fleshy cortex capable ing wide, forming a broad crown. Foliage branchlets
of water storage, enabling the species to withstand slender, usually with opposite branching, sometimes
drought spells. with more than 2 laterals from a node, spreading
horizontally at 40–80° from main axis in most cases,
1004 terete, with grooves along margins of decurrent leaf
Conservation
bases, yellowish green turning reddish to purplish
Fu & Jin (1992) determined the conservation status of brown after first year. Buds at ends of branchlets
T. jackii as Vulnerable (VU) in China Plant Red Data small; bud scales at lower node of previous year en-
Book 1, but the Conifer Specialist Group of IUCN- larged, broadly triangular, keeled, lustrous brown,
SSC (Farjon & Page, 1999) changed it to Endangered deciduous thereafter. Leaves pectinately arranged,
(EN). This species occurs very locally and has been spreading at 75–90° from shoot axis, linear to linear-
in steep decline, especially due to destruction of its lanceolate, (1.5–)2–3.5 cm long, straight or slightly
forest habitat. Special reserves to protect this species down-curved, 2.2–3.5(–4) mm wide, with 1–2 mm
in the wild are urgently needed. Natural regenera- long, twisted petiole, abruptly widening at base, ta-
tion is hampered by fragmentation of populations. pering to a long cuspidate or spinose apex, coria-
IUCN: EN (A1c) ceous, lustrous dark green on adaxial (upper) side,
smooth or with two parallel grooves at least in lower
half of blade. Stomatal bands on abaxial side only,
Uses
0.3–0.4 mm wide, yellowish, separated by a 0.5–0.8
The fragrant wood of this species is used to make mm wide, green midrib and bordered by 0.7–1 mm
agricultural implements, utensils and handicrafts; wide, flat and slightly raised green margins; stomata
outside China T. jackii is not in general cultivation small, randomly distributed. Pollen cones axillary,
and restricted to a few botanic gardens and/or pri- solitary, forming short rows on underside of lower
vate collections. leaves of lateral branchlets, with several pairs of ba-
sal bud scales in 4 rows, ca. 5–7 × 4–5 mm before
anthesis, pale yellow; microsporophylls numerous,
Torreya nucifera (L.) Siebold & Zucc., Abh. Math.- peltate, each with 4–5 marginal, pendulous pollen
Phys. Cl. Königl. Bayer. Akad. Wiss. 4 (3): 234. sacs. Seed bearing structures axillary, solitary or in
1846. Taxus nucifera L., Sp. Pl. 2: 1040. 1753. Type: pairs, sometimes grouped together on a branchlet,
Illustration: [“Fi, vulgo Kaja, Taxus nucifera”] sessile, with rounded, keeled bract scales subtending
in Kaempfer, Amoen. Exot. Fasc.: 814, 815. 1712. seed. Ripe aril including seed smooth, dark green
(lectotype). Fig. 349 ripening with purplish brown striation near apex,
obovoid to ovoid, or ellipsoid, 20–30 mm long, 13–18
Torreya ascendens Nakai ex Uyeki, [Not. Pl. Lign. mm wide, mucronate at apex. Seed proper smooth
Sikoku 11] Sci. Rep. Matsuyama Agric. Coll. 10: 3. or with 2 opposite longitudinal ridges.
1953.
Distribution
Etymology Japan: S Honshu, Kyushu, Shikoku; South Korea:
Cheju Island, Wando Island.
The species epithet (Latin nucis = nut) means ‘bear-
TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-SK
ing nuts’.
Ecology
Vernacular names
Torreya nucifera is a woodland species occurring
Japanese nutmeg tree; Kaja, Kaya (Japanese) scattered in most types of mixed broad-leaved-
conifer forest in the southern half of Japan. Wilson
Vernacular names
(1916) mentioned a locality SW of Tokyo, where it is
more abundant, growing with Abies firma on steep Florida nutmeg tree, Stinking cedar, Gopherwood
slopes composed of shale on a hill ca. 500 m high. In
other sites, including some islands in South Korea, it
Description
holds out in secondary vegetation where it becomes
shrubby. Old-growth forest with ancient trees of Small trees to 13(–18) m tall; trunk to 80 cm d.b.h.
T. nucifera in large numbers occurs in Korea’s Pija- Bark irregularly vertically fissured, sometimes
Rim National Park, with trees estimated to be 500– flaking, pale brown or greyish brown, weathering
800 years old. The altitudinal range is from near sea to dark grey. Branches spreading and ascending,
level to at least 1100 m a.s.l. Besides Abies firma and forming an open, broadly conical crown. Foliage 1005
Tsuga sieboldii, the most common conifers among branchlets slender, usually with opposite branching,
which Torreya nucifera is found, Chamaecyparis ob­ sometimes with more than 2 laterals from a node,
tusa, Podocarpus macrophyllus, Nageia nagi, Taxus spreading horizontally at 40–60° from the main
cuspidata, and Sciadopitys verticillata are often as- axis in most cases, terete, with grooves along mar-
sociated with this species. Locations dominated by gins of decurrent leaf bases, green turning yellow-
conifers in the mixed mesophytic forests of warm ish brown after first year. Buds at ends of branchlets
temperate Japan are often on rocky S-facing slopes very small; bud scales at lower node of previous year
with poorly developed soils. enlarged, broadly triangular, keeled, lustrous brown,
deciduous thereafter. Leaves pectinately arranged,
spreading at 50–80° from shoot axis, linear to linear-
Conservation
lanceolate, 1.5–3.8 cm long, straight or very slight-
IUCN: LC ly falcate, 2–3.2 mm wide, with a 0.5–1 mm long,
twisted petiole, abruptly widening at base, tapering
to a cuspidate apex, coriaceous, lustrous green on
Uses
adaxial (upper) side, slightly convex at least in lower
The wood of this species is valued in Japan for its half. Stomatal bands on abaxial side only, greyish
lustrous, pale brown leaf colour and its durability white, 0.4 mm wide, separated by a 0.6–0.8 mm
especially in contact with water. It is used for furni- wide, green midrib and bordered by 0.6–0.8 mm
ture and cabinet making, chests and boxes, Japanese wide, flat or very narrowly revolute green margins;
chessmen, and formerly water buckets. The seeds stomata small, randomly distributed. Pollen cones
are rich in edible oils and these as well as the arils are axillary, solitary, forming short rows on underside
used in Japanese cuisine. This tree is widely planted of lower leaves of lateral branchlets, with several
near temples and in parks and gardens in Japan; all pairs of basal bud scales in 4 rows, ca. 5–6 × 4.5–5
trees N of Tokyo are thought to be originally planted mm before anthesis, pale yellow; microsporophylls
(Wilson, 1916) and some of these are champion trees numerous, peltate, each with 4–6 marginal, pendu-
much larger than most trees in the forests. In Europe lous pollen sacs. Seed bearing structures axillary, in
and North America it is an uncommon ornamental pairs, sometimes grouped together on a branchlet,
shrub or tree in arboreta, botanic gardens, and oc- sessile, with rounded, keeled bract scales subtending
casionally in private gardens. seed. Ripe aril including seed smooth, dark green
streaked with purple, obovoid or broadly ellipsoid,
25–35 mm long, to 25 mm wide, mucronate at apex.
Torreya taxifolia Arn., Ann. Mag. Nat. Hist., ser. Seed proper smooth or with 2 opposite longitudinal
1, 1: 130. 1838. Type: USA: Florida, Appalachicola ridges.
River, J. Torrey s.n. (holotype K).
Distribution
Etymology
SE USA: NW Florida, SW Georgia, mainly along
The species epithet compares the leaves with those the Appalachicola River, with one population 11 km
of yew (Taxus). W of this river in Jackson Co., Florida.
TDWG codes: 78 FLA GEO
 natural range of the species. Rooted plants have now
Ecology
been shipped to gardens throughout the world in
Torreya taxifolia occurs along limestone bluffs on the hope that, if and when the natural environment
the Appalachicola River in a region with a warm and has stabilised, material from these plants can assist
humid climate, occasionally influenced in winter by the restoration of the species in its natural habitat.
cold waves from the north that dip temperatures be- Recently, “assisted migration” to areas in the south-
low the freezing point. It grows mostly in the shade ern Appalachian Mountains has been proposed,
of wooded ravines and steep, N-facing slopes under based on the hypothesis that the historical range of
canopy of Fagus grandifolia, Liriodendron tulipifera, this species in lowland Florida is a refuge from the
Acer barbatum, Liquidambar styraciflua, Quercus last Ice Age (Barlow & Martin, 2008). During the
1006 alba, and occasionally pines (Pinus taeda, P. glabra). previous interglacials it may have occurred in these
Often these woods are hung with vines (e.g. Smilax mountains (the same could be true for Taxus flori­
spp., Bignonia capreolata). Another rare conifer, dana) but, for various reasons, in the current inter-
Taxus floridana, occasionally grows with Torreya glacial it ‘missed the last bus’ and was marooned in
taxifolia. A capacity to resprout from the stem base the cool microclimate of its present location, while
probably accounts now for most of the surviving in- other trees migrated north again. It will be doomed
dividuals in the wild. if that microclimate were to disappear with ongoing
global warming. However, deliberately moving this
species north into habitats from which it would have
Conservation
disappeared naturally if it ever occurred there has
This rarest of the species of Torreya is nearly ex- also been criticized (Schwartz, 2005/08).
tinct in the wild. It is almost restricted to a 64 km IUCN: CR (A1c)
stretch of the Appalachicola River, where it occurs
sporadically in a specific limestone bluff habitat. The
Uses
number of individuals has been drastically reduced
in recent decades, probably caused by a pathological The hard and durable wood has been used for fence
fungal disease (Ascomycota) of the stem and leaves, posts locally before it became too rare and protected
first observed in 1962 and presumably alien to the under State and Federal Law. The use for Christmas
region, having arrived in the late 1950’s. Currently trees has likewise ceased. It is rare in cultivation out-
between 500 and 1000 individual plants remain, side ex situ conservation programmes aiming at re-
many only as resprouts from stumps. At the mo- introduction in the wild. This species appears to be
ment in situ conservation is considered not a viable hardy in countries with mild winters (it apparently
option; therefore, an extensive ex situ conservation can withstand occasional hard frosts). Growing this
programme has been established. This project is tree in arboreta and botanic gardens should be en-
collaborative between the Botanic Garden of Smith couraged, as it could help its conservation while be-
College and Atlanta Botanical Gardens, USA. The ing an attractive small conifer tree. It is apparently
programme has involved the collection of over commercially available from tree nurseries in South
6,000 cuttings taken from plants throughout the Carolina, USA.
Tsuga (Endl.) Carrière, Traité Gén. Conif.: 185. 1855. Type: Tsuga sieboldii Carrière
[Abies tsuga Siebold & Zucc.] (Pinaceae).

Hesperopeuce (Engelm.) Lemmon, [Pines Pacific

Distribution
Slope] Bienn. Rep. Calif. State Board Forest. 3: 126.
1890 [Tsuga sect. Hesperopeuce Engelm.]. Type: North America: (disjunct) SE Alaska, British Co-
Hesperopeuce mertensiana (Bong.) Lemmon [­Pinus lumbia, northern Rocky Mountains, Cascade Range
mertensiana Bong.]; ×Hesperotsuga C. N. Page, and Sierra Nevada of California; Great Lakes to
Notes Roy. Bot. Gard. Edinburgh 45: 389. 1989. Type: Nova Scotia, Appalachian Mountains. Asia: Sino-
×Hesperotsuga jeffreyi (A. Henry) C. N. Page [Tsuga Himalayan mountain system; Central and SE China; 1007
pattoniana (Balf.) Sénécl. var. jeffreyi A. Henry]. N Vietnam; Taiwan; Japan (disjunct).

Tsuga is the Japanese name for Hemlock (the coni-

Synopsis
fer, not the poisonous herb Conium maculatum in
Apiaceae). A total of 24 (25) species has been described in the
genus Tsuga, one of these (T. ×jeffreyi) as a notho­
species; in addition to these 1 subspecies and (exclud-
Description
ing cultivars in T. canadensis etc.) 3 varieties. Flous
Monoecious evergreen trees, usually monopodial. (1936) recognized 18 species, but generally about
Resin canals in leaves and seed cones. Bark rough 10 species are accepted by contemporary ­authors:
and scaly, longitudinally fissured in lower part of 4 in North America, 6–7 in Asia. Tsuga longibracte­
trunk. Branches plagiotropic, growing in rhyth- ata Cheng has here been treated in a distinct genus
mic pseudo-whorls, higher order branches in more Notho­tsuga, and T. mertensiana (Bong.) Carrière in a
or less horizontal or drooping sprays (Massart’s distinct section within Tsuga. Beginning with Engel-
model), sometimes with weak shoot dimorphism mann (1879) a division of the species of Tsuga in two
(T. mertensiana). Buds ovoid or globose and not or sections has been commonplace, with T. mertensi­
only slightly resinous. Leaves on spirally arranged ana (sometimes together with T. mertensiana subsp.
pulvini, usually short petiolate, broadly ligulate lin- mertensiana var. jeffreyi as a species) in one section:
ear, or narrowly ovate linear in some species, dor- Hesperopeuce Engelm., and all other species in the
siventrally flattened, entire or denticulate and with second section Tsuga (for which proposed names
an obtuse acute or truncate emarginate apex, hypo­ such as Micropeuce and Eutsuga are invalid under
stomatic or amphistomatic (T. mertensiana). Pollen the rules of the Code). This will be followed here, as
cones axillary, solitary, pedunculate, subglobular; in spite of some more or less consistent differences
microsporophylls with 2 pollen sacs containing involving entire or denticulate leaf margins, a fur-
pollen with a ring-shaped saccate structure near ther division of the genus Tsuga would in my opin-
the distal pole. Seed cones solitary, subterminal on ion rest on a too narrow basis and is unlikely to be
second year’s shoots, sessile or short pedunculate, corroborated by a phylogenetic analysis.
more or less erect at pollination but soon becom-
ing pendulous, falling without the short peduncle. Genus Tsuga (Endl.) Carrière
Bracts broadly rhombic and 1/6–1/5 the length of (Type: T. sieboldii)
the seed scales, hidden in mature cones. Seed scales Sect. Tsuga
with short, petiolate, more or less auriculate bases, Species: T. sieboldii (type), T. caroliniana,
peltate to (sub)orbicular, persistent. Seeds small, T. chinensis, T. diversifolia, T. forrestii,
covered on one side by a membranous cup, which T. canadensis, T. dumosa, T. heterophylla
extends a little over the other side and continues in Sect. Hesperopeuce Engelm.
an obliquely ovate, persistent, pale and thin seed Species: T. mertensiana
wing. Seedlings with 3–6 cotyledons.

9 species.
 nearly circular seed scales not opening wide.
Key to the sections, species and subspecies of
Leaf apex usually emarginate 7
Tsuga
7a. Leaves short, usually 0.8–1.8 cm, widest near
1a. Leaves thick, 3–4 sided or with convex sides, truncate-emarginate apex, pectinately arranged,
with stomata on both faces, mostly assurgent but shortest leaves assurgent above shoot 8
on small lateral shoots and often glaucous. Seed 7b. Leaves longer, 1–2.5 cm, with parallel margins;
cones large, ovoid-oblong to cylindrical all leaves pectinately arranged 9
Sect. Hesperopeuce 2 8a. Young shoots usually glabrous, shiny light
1b. Leaves distinctly flat, mostly with stomata on brown. Leaves irregularly arranged, with dull
the abaxial side separated by a midrib, spread- white stomatal bands. Seed cone scales in-
1008 ing or pectinately arranged, usually not assur- curved (hooded), heavily striated, buff
gent, green above, whitish below. Seed cones T. sieboldii
small (max. 4 cm long), ovoid-oblong to sub- 8b. Young shoots pubescent, orange-brown, later
globose Sect. Tsuga 3 dull brown. Leaves mostly parted and pecti-
2a. Seed cones 2–5.5 × 1.1–2.5 cm; seed scales small, nate, with niveous white stomatal bands. Seed
50–72, close together, usually dark brown cone scales flat, smooth or finely wrinkled, dark
T. mertensiana subsp. mertensiana brown T. diversifolia
2b. Seed cones 3.5–8.1 × 1.9–3.3 cm; seed scales 9a. Young shoots (remotely) pubescent. Leaves
large, 40–52, wide apart, usually light brown narrowly ovate-linear or curved-linear; mar-
T. mertensiana subsp. grandicona gins entire; apex emarginate or obtuse
3a. Margins of leaves denticulate; apex obtuse- T. forrestii
acutish, not emarginate 4 9b. Young shoots usually glabrous. Leaves linear;
3b. Margins of leaves usually entire; apex emargin- margins (of young leaves) denticulate near
ate or obtuse 6 emarginate (rarely entire) apex
4a. Leaves short, 0.5–1.8 cm long, mostly parted T. chinensis & vars.
and inversed (stomatal bands on underside);
smallest leaves not inversed (stomatal bands on
upperside) and pressed forward above shoot; Tsuga canadensis (L.) Carrière, Traité Gén. Conif.:
bands of stomata with 5–6 lines each. Seed 189. 1855. Pinus canadensis L., Sp. Pl., ed. 2, 2: 1421.
cones with acutish apex when closed 1763. Type: [“Habitat in America septentrionali”],
T. canadensis J. Clayton 547 (lectotype BM).
4b. Leaves usually longer, but variable, up to 2.5 cm
or longer, nearly all parted and inversed; bands
Etymology
of stomata with 8–10 lines each. Seed cones
with obtuse apex when closed 5 The species epithet means ‘from Canada’.
5a. Leaves narrowly ovate-linear (widest near
base), denticulate towards apex. Young branch-
Vernacular names
lets with scattered short hairs. Seed cones
broadly ovoid when closed T. dumosa Eastern hemlock, Canada hemlock
5b. Leaves narrowly elliptic, denticulate along the
whole length of margins. Young branchlets
Description
lanate pubescent with short brown hairs, mixed
with long whitish hairs. Seed cones (ovoid-) Trees to 30–40(–48) m tall, d.b.h. to 1.5–2 m; trunk
oblong when closed T. heterophylla straight, columnar, rarely forked. Bark scaly, becom-
6a. Seed cones ovoid-elliptic when closed, 2–4 cm ing rough, fissured, light cinnamon brown, turning
long; oblong seed scales opening wide or re- dark grey with age. Branches spreading mostly hori-
flexed when ripe. Leaf apex truncate or slightly zontally; branches of second order drooping at ends;
emarginate T. caroliniana crown pyramidal in young trees, with drooping
6b. Seed cones ovoid or ovoid-oblong when closed, leader, in old trees often flat topped. Branchlets pale
usually not longer than 2.5(–3) cm; (mostly) brown or grey-brown, soon grey, minutely ridged
and grooved, lanate with yellowish brown pubes- laricina, Betula spp., Acer saccharum, Quercus rubra,
cence; pulvini small, decurrent, nearly appressed Fraxinus americana, F. nigra, Fagus grandifolia, Pop­
or slightly raised. Vegetative buds conical, acute, ulus spp., and other species. It is very shade tolerant
2–3 mm long, not or slightly resinous, red-brown. and allows very little vegetation to develop under its
Leaves mostly spreading; small leaves above the own canopy.
shoot appressed, directed forward, not (!) inversed,
showing white stomatal side; other leaves more or
Conservation
less pectinate, parted below shoot, (0.5–)0.8–1.5
(–1.8) cm long, 1.5–2.2 mm wide, obliquely pedun- IUCN: LC
culate and twisted at base, ligulate-linear or widest
near base, denticulate at margins, grooved above, 1009
Uses
flattened, obtuse at apex; stomata in two white bands
on the abaxial (lower) side; leaf colour dark green The slow growing Eastern hemlock produces lum-
on upperside. Pollen cones numerous, 3–5 mm long, ber of good quality suitable for building (e.g. roofs,
(orange-)yellow. Seed cones numerous on all outer floors) and making crates or boxes, but until recently
branchlets, on delicate, pubescent, 4–5 mm long these kinds of use were completely overshadowed by
peduncles, ovoid-cylindrical with acute apex when its use in the paper pulp industry. Other former uses
closed, ovoid and obtuse when opened, 1.5–2.5(–3) were to make telegraph poles and railway sleepers.
cm long, 1–1.8(–2.2) cm wide with opened scales, In the past its bark was used in the tanning industry.
ripening to light brown or grey-brown. Seed scales Eastern hemlock is still in demand as an ornamental
obovate-orbicular, convex, thin and papery (cones tree; it was introduced to Europe in 1736. In gardens
very light), 10–12 × 6–8 mm at mid-cone; exposed and parks it often grows several trunks, but this is
abaxial surface striated or wrinkled, glabrous, with by no means a characteristic of the species in its nat-
impressions of two lower scales in lower part; up- ural habitat. A large number of cultivars has been
per margin entire or obscurely denticulate, often produced, including variegated foliage plants and
undulate; base narrowed, short pedicellate. Bracts dwarf forms raised from cuttings, whereby the slow
3–4 mm long, red-brown. Seeds ovoid-oblong or growth of this species is an obvious advantage over
nearly reniform, 3 mm long, light brown; seed wings Western hemlock, of which few cultivars exist. Con-
ovate or ovate-triangular, 6–7 × 4 mm, pale yellow- versely, due to that slower growth Eastern hemlock
ish transparent. is less preferred as a forestry plantation tree, giving
way to Western hemlock.
Distribution
E North America: from Nova Scotia to N Georgia, Tsuga caroliniana Engelm., Bot. Gaz. 6: 223. 1881.
westwards to Minnesota. Type: USA: South Carolina, A. H. Curtiss s.n. [ex
TDWG codes: 72 NBR NSC ONT PEI QUE 74 MIN herb. Engelmann] (lectotype MO).
WIS 75 CNT INI MAI MAS MIC NWH NWJ NWY OHI
PEN RHO VER WVA 76 GEO KTY NCA SCA TEN VRG
Etymology
The species epithet refers to the States of North and
Ecology
South Carolina in the USA.
Tsuga canadensis occurs from near sea level (Nova
Scotia) to 600 m in N Michigan, in the southern
Vernacular names
Appalachians between 600 and 1500 m a.s.l. The
soils are of glacial, fluvio glacial, alluvial, or colluvial Carolina hemlock
origin, podzolic and usually highly acidic (pH 3–4).
The climate is cool and humid, with annual precipi-
Description
tation between 700 and 1500 mm. Tsuga canadensis
grows locally pure, but is usually mixed with other Trees to 20–25 m tall; trunk to 50–60 cm d.b.h.,
conifers and broad-leaved trees: Pinus strobus, P. res­ straight, columnar. Bark becoming rough and scaly,
inosa, Abies balsamea, Picea rubens, P. glauca, Larix fissured, with purplish grey outer bark and red-
 brown inner bark exposed in the fissures. Branches sons. It is a rare and scattered tree, which grows sin-
spreading horizontally, but lower branches curved gly, mixed with broad-leaved trees and shrubs, or in
downward; branches of second order drooping at small nearly pure groves of only a few individuals.
ends; crown of young trees conical, with droop-
ing leader, in old trees broader, open or dense and
Conservation
flat-topped. Branchlets red-brown, paler orange-
brown below, shiny, with fine grooves and ridges This species has a sufficiently large range (extent of
and short, dark brown pubescence especially in the occurrence, EOO), but it occurs within that range
grooves; pulvini well developed, swollen, appressed only in scattered, small sub-populations at cool,
and decurrent. Vegetative buds ovoid-conical, 3–4 moist or rocky sites where most other (angiosperm)
1010 × 1.5–2.5 mm, not or only slightly resinous, red- trees and shrubs thrive less well. It could be at risk
brown. Leaves pectinate at nearly right angles from from succession by these angiosperms, if climate
shoot, remote, a few above shoot directed forward, change were to affect these localities trending to-
of unequal length without a ranked order, 0.5–2 cm wards warmer and drier conditions in future.
long, 1.8–2 mm wide, linear, with entire margins, IUCN: NT
flattened, grooved above, truncate, retuse or slightly
emarginate at apex; stomata in two white bands on
Uses
the underside (abaxial surface); leaf colour on up-
perside dark, lustrous green. Pollen cones ca. 5 mm The wood of Carolina hemlock has similar proper-
long, yellow tinged with purple. Seed cones on 4–5 ties as that of Eastern hemlock, but the species is
mm long, pubescent peduncles, ovoid-elliptical much less common and widespread, so its commer-
when closed, with opened scales irregular, more or cial value is limited. As an ornamental tree it is also
less ovoid, 2–3.5(–4) cm long, 1.5–2.5(–3) cm wide, less commonly used, being somewhat more difficult
maturing to light brown or red-brown. Seed scales to establish from seed and growing rather slowly.
ovate-oblong to oblong, opening at right angles to A few cultivars have been selected, mostly of com-
rachis or recurved, 13–20 × 6–10 mm, abaxial sur- pact or dwarfish growth habits.
face striated on exposed part, slightly puberulent
when green, but soon glabrous; adaxial side with
two dark seed wing marks; upper margin entire, Tsuga chinensis (Franch.) E. Pritz., Bot. Jahrb. Syst.
rounded, obtuse or tapering, sometimes undulate; 29: 217. 1900.
base short pedicellate. Bracts 4–5 mm long, weakly
trilobate, pale brown. Seeds ovoid-cuneate, 3–4.5 ×
Etymology
1.5–2.5 mm, light brown with dark spots; seed wings
oblong, tapering to apex, 8–12 × 3.5–5 mm, light yel- The species epithet refers to China.
lowish brown, transparent.
Vernacular names
Distribution
Chinese hemlock; Tieshan (Chinese)
USA: Appalachian Mountains.
TDWG codes: 78 GEO NCA SCA TEN VRG
Description
Trees to 40–50 tall, d.b.h. to 1.5–2 m; trunk mono­
Ecology
podial, often forked above 1/2 height. Bark on trunk
Tsuga caroliniana occurs in low to medium high rough and scaly, with numerous brown-grey plates.
mountains, at altitudes between 600 m and 1500 m Branches assurgent, spreading more horizontal-
a.s.l. (commonly 750 m to 1200 m), on rocky, moist ly at ends; crown broad, conical, or flat topped in
N- or E-facing slopes or rocky ridges, also along old trees. Branchlets pale yellowish brown, finely
streams in cool ravines. The climate is humid and grooved between slightly swollen, appressed, decur-
cool, relatively warm at lower elevations, the annual rent and darker pulvini, with minute pubescence
precipitation exceeds 1000 mm and falls throughout in grooves, soon glabrous. Vegetative buds ovoid-
the year; there is much cloudy weather in all sea- globose, 1–4 mm long, not resinous, dark brown
or red-brown. Leaves mostly pectinate, but a few the country. The wood is hard and durable and used
short leaves erect, (0.6–)1–2(–2.7) cm long, 1.8–3 for construction, shingles for roofing, general car-
mm wide, (broad) linear, flattened, grooved above, pentry, and joinery. This species was introduced by
with near apical margins of young leaves denticu- Ernest Wilson for the Veitch Nurseries in England
late, emarginate, sometimes entire at apex; stomata in 1900, but it has remained uncommon in cultiva-
in two white bands on underside (abaxial side) sepa- tion. No cultivars have been recorded. In plantation
rated by a midrib, leaf colour green on upperside. forestry, it is increasingly planted in the eastern USA
Pollen cones crowded near ends of branchlets, 3–5 as a substitute for T. canadensis and T. caroliniana,
mm long, yellow, with purple tinge. Seed cones nu- because it is resistant to an insect pest that adversely
merous, short pedunculate or sessile, ovoid-oblong affects the native species.
when closed, subglobular or ovoid-oblong when 1011
opened, 1.5–3(–4?) cm long, 1.3–2.2(–3.5?) cm wide, 3 varieties are recognized:
light green when immature, ripening to light, glossy
brown. Seed scales nearly circular, convex, 8–12 Tsuga chinensis (Franch.) E. Pritz. var. chinensis.
(–14) × 8–10(–12) mm; abaxial surface usually stri- Abies chinensis Franch., J. Bot. (Morot) 13 (8):
ated, shining, with imprints of two lower scales, gla- 259. 1899. Type: China: Chongqing Municipality,
brous; upper margin rounded, truncate or retuse; Chengkou Xian, Daba Shan, P. G. Farges 808
base short pedicellate. Bracts transversely rhombic, (holotype P). Fig. 350, 351
with denticulate upper margin, 1–2(–3) mm long.
Seeds ovoid-oblong or ovoid-triangular, 3–4 × 2 Tsuga formosana Hayata, Gard. Chron., ser. 3, 43:
mm, light brown; seed wings obliquely ovate, 6–7 × 194. 1908; Tsuga chinensis (Franch.) E. Pritz. var. for­
3.5 mm, light yellowish, transparent. mosana (Hayata) H. L. Li & H. Keng, Taiwania 5: 64.
1954.
Tsuga patens Downie, Notes Roy. Bot. Gard. Edin-
Distribution
burgh 14: 16. 1923; Tsuga chinensis (Franch.) E. Pritz.
S and N Central and SE China, SE Xizang [Tibet]; subsp. patens (Downie) E. Murray, Kalmia 12: 26.
Taiwan. 1982; Tsuga chinensis (Franch.) E. Pritz. var. patens
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- (Downie) L. K. Fu & Nan Li, Novon 7 (3): 263. 1997.
SC CHC-YN CHN-GS CHN-SA CHS-AH CHS-FJ CHS- Tsuga tchekiangensis Flous, Bull. Soc. Hist. Nat.
GD CHS-GX CHS-HE CHS-HN CHS-JX CHS-ZJ CHT Toulouse 69: 414. 1936; Tsuga chinensis (Franch.)
38 TAI E. Pritz. var. tchekiangensis (Flous) W. C. Cheng &
L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 119. 1978.
Tsuga chinensis (Franch.) E. Pritz. var. daibuensis
Ecology
S. S. Ying, Bull. Exp. Forest Natl. Taiwan Univ. 114:
Tsuga chinensis and its varieties occur at altitudes 150. 1974.
between (600–)1200–3200 m a.s.l. The soils are red
and yellow earth, or mountain podzols at high el-
Description
evations. The climate is cool temperate, moist (an-
nual precipitation 1000 mm to 2000 mm) or very Seed cones ovoid-oblong when closed, more or less
wet (Taiwan). This species is widely distributed and subglobose when opened, 1.5–2.5 cm long, 1.3–2.2
occurs in the mixed mesophytic forest formation cm wide; seed scales nearly circular, short petiolate,
(Wang, 1961) together with numerous broad-leaved 8–12 × 8–10 mm.
trees and several conifers; on the Southwestern Pla-
teau also in the montane coniferous forests with
Distribution
Abies, Picea and other conifers.
As for the species.
Uses
Conservation
Chinese hemlock is a valuable timber tree in China
and has been logged extensively in many parts of IUCN: LC
 Tsuga chinensis (Franch.) E. Pritz. var. oblongi­
Etymology
squamata W. C. Cheng & L. K. Fu, Acta Phyto-
tax. Sin. 13 (4): 83. 1975. Tsuga oblongisquamata The species epithet refers to the the different lengths
(W. C. Cheng & L. K. Fu) L. K. Fu & Nan Li, Novon of leaves on a shoot.
7 (3): 263. 1997. Type: China: Hubei, W Hubei, Chi­
nese collector 950 (holotype PE).
Vernacular names
Northern Japanese hemlock; Kome-tsuga, Kuro-
Description
tsuga (Japanese)
Seed cones ovoid-oblong when opened, 1.8–2.8 ×
1012 1–1.5 cm; seed scales obovate-oblong, distinctly peti-
Description
olate, 10–14 × 6–9 mm.
Trees to 20–25 m tall, d.b.h. to 50–60 cm; trunk
Distribution straight or curved, often forked above half of
height. Bark on trunk rough and scaly, longitudi-
China: Gansu (Zhouqu), W Hubei, Sichuan.
nally grooved, dark brown-grey. Branches spreading
TDWG codes: 36 CHC-HU CHC-SC CHN-GS
horizontally or ascending in top; crown of mature
trees broad, domed and dense, more open in old
Conservation
trees. Branchlets orange-brown, later dull brown,
IUCN: VU [B2 a-b(iii)] ridged and grooved between appressed, decurrent
pulvini, short pubescent, soon glabrous. Vegetative
Tsuga chinensis (Franch.) E. Pritz. var. robusta buds obovoid or globular, 2–3 × 2–2.5 mm, not or
W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): only slightly resinous, dark brown. Leaves pectinate
83. 1975. Type: China: Hubei, Yangtse Gorges, [?] or at least parted, but shorter leaves above shoot
Chen et al. 2000 (holotype PE). assurgent, (0.5–)0.8–1.5(–1.8) cm long, ca. 2 mm
wide, ligulate-linear, slightly wider near truncate-
emarginate apex, slightly curved or straight, flat-
Description
tened, grooved above; stomata in two broad, white
Seed cones ovoid-oblong when closed, more or less bands separated by a midrib on the abaxial (under)
subglobose when opened, larger than in var. chin­ side; leaf colour dark lustrous green on upperside.
ensis (possibly up to 4 × 3.5 cm); seed scales robust Pollen cones 3–5 mm long, yellow when shedding
and thick. pollen. Seed cones numerous on outer branchlets
of the entire crown, on 3–4 mm long, pubescent
Distribution peduncles, ovoid when closed, subglobose when
opened, (1.5–)2–2.5 cm long, 1.5–2 cm wide, green
China: Hubei, Sichuan (Yalong Valley).
or purplish green, ripening to shiny, reddish brown
TDWG codes: 36 CHC-HU CHC-SC
or dark brown. Seed scales nearly circular to obo-
vate, flat or slightly convex, spreading at ca. 90° from
Conservation
rachis, 8–12 × 7–10 mm; abaxial surface smooth or
IUCN: VU [B2 a-b(iii)] finely wrinkled, with imprints of two overlapping
scales; base broad cuneate, sometimes short pedi-
cellate. Bracts broad, truncate, with two small api-
Tsuga diversifolia (Maxim.) Mast., J. Linn. Soc., cal teeth, 2–3 mm long. Seeds ovoid, ca. 3 × 2 mm,
Bot. 18: 514. 1881. Abies diversifolia Maxim., Bull. (dark) brown; seed wings obliquely ovate, 6–8 × 3–4
Acad. Imp. Sci. Saint-Pétersbourg 12: 229. 1868. mm, yellowish or orange-brown, transparent.
Type: not designated.
Distribution
Tsuga blaringhemii Flous, Bull. Soc. Hist. Nat. Tou-
louse 69: 410. 1936; Tsuga diversifolia (Maxim.) Japan: Honshu, Kyushu.
Mast. subsp. blaringhemii (Flous) E. Murray, Kalmia TDWG codes: 38 JAP-HN JAP-KY
12: 26. 1982.
 Jahrb. Syst. 29: 217. 1900; Tsuga dumosa (D. Don)
Ecology
Eichler var. yunnanensis (Franch.) Silba, Phytologia
Tsuga diversifolia occurs in the mountains at al- 68: 73. 1990.
titudes between 700 m and 2000 m a.s.l., on usu- Tsuga dura Downie, Notes Roy. Bot. Gard. Edin-
ally podzolic soils developed on volcanic or igneous burgh 14: 16. 1923; Tsuga yunnanensis (Franch.)
rock. The climate is cool, with cold, snowy winters E. Pritz. subsp. dura (Downie) E. Murray, Kalmia 12:
and abundant rainfall in summer (annual precipita- 26. 1982.
tion 1000 mm to 2500 mm). It is in many areas the Tsuga wardii Downie, Notes Roy. Bot. Gard. Edin-
most common tree species in mixed coniferous for- burgh 14: 17. 1923; Tsuga chinensis (Franch.) E. Pritz.
ests, being very shade tolerant. Other common coni- subsp. wardii (Downie) E. Murray, Kalmia 12: 26.
fers are Picea jezoensis, Abies homolepis, A. veitchii, 1982. 1013
A. mariesii (at high elevations), Larix kaempferi, Pi­ Tsuga leptophylla Hand.-Mazz., Akad. Wiss. Wien,
nus parviflora, Thuja standishii, and Thujopis dola­ Math.-Naturwiss. Kl., Anz. 61: 83. 1924; Tsuga du­
brata var. hondae; broad-leaved trees are e.g. Betula mosa (D. Don) Eichler subsp. leptophylla (Hand.-
ermanii, B. corylifolia, Sorbus japonica, Alnus hirsuta Mazz.) E. Murray, Kalmia 12: 26. 1982.
var. sibirica, and Quercus mongolica var. grosseserra­
ta. Rhododendron spp. and/or Sasa spp. may form a
Etymology
dense undergrowth in the shrub layer, in other, very
wet areas only thick moss layers carpet fallen logs ’Dumosa’ is a Japanese name for Tsuga (but not re-
and the forest floor. ferring to this species).

Conservation Vernacular names

IUCN: LC Himalayan hemlock; sula, thingia (Nepalese); tang-
shing (Bhutan); Yunnan tieshan (Chinese)
Uses
Description
Northern Japanese hemlock is exploited in Japan for
timber. In the past it mainly provided pulp for paper, Trees to 40–50 m tall, d.b.h. to 1.5–2.5(–2.7) m; trunk
but now this commodity, requiring vast resources, monopodial, or multistemmed. Bark soon rough
mainly comes from abroad. Instead, the use of this and scaly, deeply fissured, with large plates in old
species has largely shifted to construction, carpentry, trees, pink-brown; outer bark grey. Branches ascend-
and joinery and, as the wood is generally dense and ing or nearly erect near tree top, spreading horizon-
moderately hard and sometimes attractively figured tally or downward below; branches of second order
with reddish brown heartwood and lighter sapwood, drooping at ends, forming flat, slanting planes of
it is used for furniture. As and ornamental tree it is foliage; crown broad conical in young trees, becom-
planted in Japanese gardens and parks; it is also used ing flat topped and irregular in old trees. Branch-
in bonsai culture. In Europe and North America it is lets pale pinkish brown to brown, later grey, ridged
less commonly used, being slow growing. A dwarf and grooved between appressed, decurrent pulvini,
form (cultivar) which grows very slow is used for with scattered pubescence. Vegetative buds obovoid
rockeries. Hemlocks are unsuitable as Christmas to globular, 2–2.5 mm long, not resinous, brown.
trees, because when cut and taken indoors they Leaves directed forward, rigid, spreading irregu-
loose their leaves sooner than any other conifer. larly, more pectinate below shoot, (1–)1.5–2.5(–3.5)
cm long, 1.5–3 mm wide, very narrowly ovate-linear,
widest near base, straight or slightly curved, flat-
Tsuga dumosa (D. Don) Eichler, in Engler & Prantl,
tened; margins (mostly on the distal half) denticu-
Nat. Pflanzenfam. 2 (1): 80. 1887. Pinus dumosa
late, but in older leaves becoming entire, grooved
D. Don, in Lambert, Descr. Pinus 2: 55. 1824. Type:
above, obtuse acutish at apex; stomata in 2 whitish
not designated.
bands separated by a midrib on abaxial (under)
Abies yunnanensis Franch., J. Bot. (Morot) 13 (8): side; leaf colour green or glaucous green on upper-
258. 1899; Tsuga yunnanensis (Franch.) E. Pritz., Bot. side. Pollen cones numerous, 3–5 mm long, yellow at
 maturity. Seed cones numerous on outer branchlets the bark these are traditionally used in the roofing
of entire crown, very short pedunculate or nearly of wooden houses. The foliage is sometimes burnt
sessile, broadly ovoid, more globular when opened, as incense in Buddhist religious shrines. This spe-
(1.5–)2–3 cm long, 1.5–2.5(–3) cm wide, light green cies has been introduced in Europe (England) in
or purplish green to purple, ripening to lustrous 1838, but is not common in cultivation; sometimes
light brown, but unexposed part of seed scales dark trees are listed under its synonym T. yunnanensis
purplish brown. Seed scales broadly ovate-elliptic, when coming from the Chinese part of its range. Its
convex, 11–14 × 9–12 mm at mid-cone; abaxial sur- planting is usually limited to arboreta and botanic
face smooth, finely striated, with imprints of over- gardens with living collections of conifers in regions
lapping scales, glabrous; upper margin rounded, with mild winters and abundant rainfall.
1014 entire, slightly recurved or straight; base petiolate,
slightly auriculate. Bracts broadly angular-ovate, ca.
3 mm long. Seeds ovoid-oblong, 3–4 × 2–2.5 mm, Tsuga forrestii Downie, Notes Roy. Bot. Gard.
brown; seed wings obliquely ovate, 6–8 × 3.5–4 mm, Edinburgh 14: 18. 1923. Tsuga chinensis (Franch.)
light yellowish brown, transparent. E. Pritz. var. forrestii (Downie) Silba, Phytologia
68: 72. 1990. Type: China: Yunnan, Lijiang Shan,
G. Forrest 17169 (holotype E). Fig. 352
Distribution
Himalaya; China: mountains of SE Xizang [Tibet],
Etymology
NW Yunnan and SW Sichuan; N Myanmar [Bur-
ma]. The species epithet commemorates the British plant
TDWG codes: 36 CHC-SC CHC-YN CHT 40 EHM- collector George Forrest (1873–1932).
AP EHM-BH EHM-DJ EHM-SI NEP WHM-UT 41 MYA
Vernacular names
Ecology
Forrest’s hemlock; li jiang tie shan (Chinese)
Tsuga dumosa occurs in the Himalaya in a belt be-
tween 2600 m and 3200 m a.s.l., in a wide range of
Description
habitats, usually on alpine lithosols. In China it is
most common between 2200 m and 2800 m a.s.l., but Trees to 25–30 m tall, d.b.h. to 1 m; trunk straight,
it occurs as low as 1700 m and up to 3500 m a.s.l. in often forked or multistemmed above half of height.
Sichuan and Yunnan. The climate is moist monsoon, Bark soon rough and scaly, orange-brown, in old
with abundant precipitation, wettest in the eastern trees fissured below and brownish grey. Branches as-
Himalayas and Upper Burma, where it can receive cending, then spreading more horizontally; branch-
up to 10,000 mm rain per year. It is an almost con- es of second order drooping at ends; crown broadly
stant companion of conifers, e.g. Abies spp., ­Picea conical in young trees, becoming domed, flat topped,
spp.; Cedrus deodara in the western Himalayas, and or irregular in old trees. Branchlets (pale) reddish
Larix griffithii in the eastern Himalayas; it is espe- brown or pink-brown, turning grey-brown, ridged
cially abundant on slopes with a northerly exposure, and grooved between appressed, apically thickened,
where it is the most shade tolerant tree. decurrent pulvini, (remotely) pubescent, soon gla-
brous. Vegetative buds globular, 2–3 mm diam., not
or slightly resinous, brown. Leaves irregularly pecti-
Conservation
nate, spreading at nearly 90° from shoot, but short-
IUCN: LC est leaves more or less erect above shoot, 1–2.5 cm
long, ca 2 mm wide, narrowly ovate-linear, straight
or curved, flattened, weakly grooved on upper-
Uses
side; margins entire, emarginate or obtuse at apex;
Himalayan hemlock is a timber tree of some impor- stomata in two glaucous white bands separated by
tance locally, but considered by Indian foresters to a prominent midrib; leaf colour lustrous green or
be inferior to several other Himalayan conifers. Its glaucous on upperside. Pollen cones 3–5 mm long,
wood can be split into shingles and together with yellow when shedding pollen. Seed cones numerous
on outer branchlets of entire crown, short pedun-
Conservation
culate or sessile, ovoid-oblong, becoming ovoid-
globose when opened, (1.6–)2–3(–4?) cm long, This species has a limited range separated into two
1.3–2(–3?) cm wide, green or greenish purple, ripen- or three disjunct areas. Deforestation and logging
ing to light brown. Seed scales nearly circular to very have substantially reduced the area of occupancy
broadly elliptic, slightly convex, 10–13 × 8–11 mm at (AOO) of this species; however, the assessment dates
mid-cone; abaxial surface finely striated, glabrous, from before 2001 when the criteria here cited were
lustrous; upper margin rounded, entire, or erose changed and now would require a decline of 50% or
in old cones; base short pedicellate. Bracts broadly more over three generations instead of a mere 20%
ovate, with denticulate margins, acutish, 3–4 mm under the 1994 criteria. A reassessment would there-
long. Seeds ovoid-oblong, 3.5 × 2 mm, brown; seed fore necessitate a new estimate of decline. 1015
wings obliquely ovate, 8 × 4 mm, yellowish brown, IUCN: VU (A1cd)
transparent.
Uses
Taxonomic notes
Forrest’s hemlock is a timber tree used for construc-
This species is very similar to T. dumosa and pos- tion, aircraft, furniture and as props for mines. This
sibly only a subspecies of it. Its characters place it species is present in several arboreta in Europe and
between T. dumosa and T. chinensis; it occurs in the North America, almost exclusively from early 20th
area where the ranges of these two species meet: century introductions made by the famous ‘plant
the Lijiang Shan and mountains to the northwest hunters’ of the time. These planted trees usually
(NW Yunnan) and north (SW Sichuan). Cheng & bear a good crop of seed cones with viable seed
Fu (1978) reported seed cones of max. 4 × 3 cm, but and in sufficiently wet places like western Scot-
neither the type (G. Forrest 17169, holo. E, iso. K), land’s arboreta, they can be seen to self-propagate
nor other collections I have seen have such large seedlings. However, since this species is rarely the
cones. Perhaps trees with such cones are T. chinensis only one of it’s genus planted there, we cannot be
var. robusta. In Flora of China 4: 41 (1999), T. for­ sure, without specialized genetic research, that the
restii has been treated as a variety of T. chinensis. seedlings are ‘pure’ T. forrestii. This creates a prob-
lem for the maintenance of the species in cultiva-
tion. If taken from cuttings, we base its propagation
Distribution
on a very narrow genetic basis; from seeds we may
China: NE Guizhou (Jiangkou: Fanjing Shan), SW introduce ‘alien’ genes into the stock of young trees.
Sichuan, NW Yunnan. Ideally, conifers like this are continually introduced
TDWG codes: 36 CHC-SC CHC-YN from correctly identified natural sources. Under the
current restrictive legislation pertaining to plant
collecting and international traffic of the same and
Ecology
its propagules, this option has become much more
Like the other two species occurring on the SW Pla- difficult to realize. In the long term, ill-considered
teau of China, T. forrestii is a high mountain spe- legislation may seriously hamper the perpetuation
cies occurring between 2000 m and 3500 m a.s.l. of laboriously assembled species collections in arbo-
The soils are mostly podzolized. The climate is tem- reta and botanic gardens.
perate to cold temperate, with annual precipitation
between 1000 mm and 2000 mm. It is a constitu-
ent of the montane boreal coniferous forest forma- Tsuga heterophylla (Raf.) Sarg., Silva N. Amer. 12:
tion, where it is mixed with Abies spp., Picea spp., 73, t. 605. 1899. Abies heterophylla Raf., Atlantic J. 1:
Larix potaninii, occasionally Pseudotsuga sinensis, 119. 1832. Type: not designated. Fig. 353
Cephalo­taxus fortunei, and broad-leaved trees, e.g.
Betula albosinensis, Acer spp., Sorbus spp. Quercus
Etymology
spp., and Magnolia spp. Tsuga forrestii remains in
most places a minor component of the forest (Wang, The species epithet describes the variable size of the
1961). leaves on a single shoot.
 Vernacular names Ecology
Western hemlock, Pacific hemlock Tsuga heterophylla occurs from sea level to 600 m
a.s.l. along the Pacific coast, in the Rocky Moun-
tains it reaches to 1800 m. It grows on a variety of
Description
soils with an acid organic top layer (pH 3.5–5). The
Trees to 60–70 m tall, d.b.h. to 1.5–2.5 m; trunk climate is cool maritime along the coast, cold mon-
straight, often more or less buttressed at base (from tane in the interior, the annual precipitation varies
originating on a ‘nurse log’). Bark on trunk rough between (500–)900–3800 mm, decreasing towards
and scaly, fissured below, dark grey. Branches spread- the interior. Dry summers limit its range in the
1016 ing horizontally, drooping at ends; crown in young Rocky Mountains. It is highly sympatric with ­Picea
trees conical, with drooping leader, in old trees sitchensis in most of the range. It is extremely shade
broad conical. Branchlets reddish brown above, pale tolerant, but has a shorter life span than Pseudo­
yellowish below, turning grey in second year, finely tsuga menziesii or Picea sitchensis. Close along the
ridged and grooved between small, dark tipped, ap- coast it may form occasionally pure stands, but
pressed, decurrent pulvini, lanate pubescent with more commonly it is an important constituent of
short brown hairs, mixed with fewer long whitish the (maritime) mesothermal coniferous forest. On
hairs, glabrescent after third year. Vegetative buds the Olympic Peninsula of Washington it reaches
ovoid-conical, 1.5–2 mm long, not resinous, densely maximum size, together with other giant conifers.
pubescent, pale brown or reddish brown. Leaves of Its tolerance to shade allows it to grow up under the
unequal length, with longer leaves spreading side- canopy of other trees, but a thick moss layer usu-
ways or radially, parted below the shoot, 0.7–2(–2.3) ally prevents the light seeds from reaching the soil.
cm long, 1.5–2 mm wide, (very) narrowly elliptic, Instead, seeds germinate massively on fallen trees
flattened, grooved above; margins denticulate; apex (‘nurse logs’), from where a few saplings are able to
obtuse or slightly emarginate; stomata in 2 white send roots down into the soil; as a result T. hetero­
bands on abaxial (under) side separated by a midrib; phylla often stands in rows (‘collonades’) long after
leaf colour dark green on upperside. Pollen cones the nurse log has rotten away.
subglobular, 3–5 mm long, crimson or light red, yel-
low at anthesis. Seed cones numerous on all outer
Conservation
branchlets, on 4–6 mm long, pubescent peduncles,
ovoid-oblong when closed, subglobular with opened IUCN: LC
seed scales, 1.4–2.2(–3) cm long, 1.5–2.5 cm wide,
light green or grey-green, occasionally tinged violet,
Uses
ripening to light brown, but unexposed part of seed
scales turning dark brown. Seed scales suborbicular, Western hemlock is an important timber tree in
ovoid-oblong or oblong, thin, opening wide, 6–13 × the Pacific Northwest (USA) and W Canada. Un-
5–9 mm at mid-cone; exposed part of abaxial sur- like its eastern sister species it is a fast grower even
face striated or wrinkled, glabrous; upper margin outperforming Douglas fir. The timber is used for
rounded or obtuse; base short pedicellate. Bracts pilings, poles and railway sleepers and especially
triangular, 2–4 mm long. Seeds ovoid-oblong, 2–3.5 for construction. A large proportion of the annual
mm long, (light) brown; seed wings ovate triangular, harvest goes to the wood pulp industry for various
4–10 mm long, light yellowish brown, transparent. applications. This species has been introduced in
Britain and other parts of NW Europe as a forest-
ry plantation tree; in the wetter parts near the At-
Distribution
lantic coast it will regenerate spontaneously. As an
W North America: along the coast from Alaska to extremely shade tolerant conifer it is suitable as an
N California and in the Cascade Range, also in the under planted tree in deciduous broad-leaved forest;
northern Rocky Mountains (mainly in British Co- however, as it in turn shades out every growth under
lumbia and Idahoe). it, this should not be done in (semi) natural wood-
TDWG codes: 70 ASK 71 ABT BRC 73 IDA MNT ORE land where the indigenous flora is valued. Although
WAS 76 CAL frequently planted as a specimen tree in arboreta
and parks, mainly within its natural range and in the × 2–2.5 mm, brown; seed wings oval-oblong, 4–7 ×
British Isles, this species is not much used in horti- 3–3.5 mm, light brown.
culture and few cultivars have been raised.
Distribution
Tsuga mertensiana (Bong.) Carrière, Traité Gén. Pacific Coast Region of NW North America: from
Conif., ed. 2, 1: 250. 1867. Alaska to California, in the Cascade Range and Sier-
ra Nevada, and isolated occurrences in the northern
Rocky Mountains.
Etymology
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
This species was named after Karl Heinrich Mertens 1017
(1796–1830), whose plant collections from the
Ecology
American Pacific coast were studied by A. H. von
Bongard in St. Petersburg. Tsuga mertensiana is a subalpine species, occurring
from near sea level in Alaska to 1500 m a.s.l. along
the coast; in the Cascade Range between 1200 m and
Vernacular names
2100 m a.s.l., and in the Sierra Nevada (subsp. gran­
Mountain hemlock dicona) between 1800 m and 3350 m a.s.l. It grows on
a variety of non-alcareous acidic soils, sometimes on
peat, more commonly on mor humus (pH 3.1–3.9).
Description
Subsp. mertensiana is restricted to a climatic zone
Trees to 30–40(–45) m tall, d.b.h. to 1–1.5 m; trunk with high precipitation, in British Columbia between
straight, but often curved or almost prostrate at tree 2000 mm and 4000 mm per year, with long, snowy
limit. Bark deeply fissured in the lower part of the winters and short, cool summers. Subsp. grandicona
trunk, dark reddish brown. Branches spreading or grows in a much drier climate, but there primari-
assurgent; crown usually narrowly conical, but often ly on high, N-facing slopes. The species is a major
deformed by wind. Branchlets yellowish or orange- component of the Mountain hemlock-Subalpine fir
brown, turning grey in 2–4 years, with ridges end- forest, occurring in pure stands or mixed with Abies
ing in decurrent pulvini with oval leaf scars, densely lasiocarpa, locally also with A. amabilis, Picea glau­
yellowish pubescent. Vegetative buds ovoid conical, ca, P. sitchensis, P. engelmannii (Rocky Mts.), Pinus
2–3 × 1–2 mm, not resinous, brown. Leaves densely spp., Tsuga heterophylla, Xanthocyparis nootkatensis,
covering shoot, directed forward, ascending above Juniperus occidentalis, and Betula papyrifera.
shoot, (0.5–)0.7–2(–2.5) cm long, 1–1.5 mm wide,
short petiolate at base, linear, straight or curved,
Uses
thick, 3–4 sided or with convex sides, shallowly
grooved near base, obtuse or acutish at apex; sto- This slow growing species produces moderately
mata in several lines on all sides; leaf colour green strong wood, but its use is limited due to environ-
or glaucous green. Pollen cones pendulous, ca. 1 cm mental considerations. It is soft and close-grained,
long, at first purplish blue, then yellow with purple with brown heartwood, sometimes pinkish, and
tinge. Seed cones mostly near top of tree, erect at lighter sapwood; its uses are now restricted to car-
first, later mostly pendant, short pedunculate or pentry and some limited construction applications.
sessile, ovoid-oblong to cylindrical, obtuse at apex, In its natural habitat it is much appreciated by hikers
(2–)3–5.5(–8.1) cm long, (1.1–)1.5–2.5(–3.3) cm wide for its picturesque appearance on mountain ridges.
with opened scales, purplish blue when young, rip- It also makes an excellent ornamental tree for gar-
ening to dark brown or light brown. Seed scales 40– dens with its dense foliage growing from long and
72(–80?), obovate-cuneate, 10–13(–20) × 7–10(–15) short shoots and naturally conical habit. Despite
mm at mid-cone, spreading very wide or reflexed this, it is uncommon in gardens and only a limit-
when mature, puberulent when immature, but soon ed number of cultivars is known. Among these the
glabrous; upper margin rounded, entire; base short ‘blue’ or glaucous leaf forms or selections are espe-
pedicellate. Bracts ligulate-cuspidate, 4–7 mm long, cially valued. High altitude provenances may be sus-
visible with opened seed scales. Seeds cuneate, 3–5 ceptible to ‘late’ frost, but those from more coastal
 northern areas should not suffer from this kind of 2005). It was at some time considered to be a hybrid
damage as much. between T. heterophylla and T. mertensiana, but no
such hybrids have been confirmed to occur in na-
2 subspecies and 2 varieties are recognized:
ture by recent investigators.
Tsuga mertensiana (Bong.) Carrière subsp. merten­
siana var. mertensiana. Pinus mertensiana Bong., Distribution
Mém. Acad. Imp. Sci. Saint-Pétersbourg, sér. 6,
Canada: British Columbia (Vancouver Island).
Sci. Math. 2: 163. 1832; Hesperopeuce mertensiana
TDWG codes: 71 BRC
(Bong.) Rydb., Bull. Torrey Bot. Club 39: 100. 1912.
Type: not designated. Fig. 354
1018
Ecology
Description
Presumably mixed coniferous forests in the Pacific
Short shoots apparent. Leaves usually crowded and Coast Region of North America.
ascending above shoots. Seed cones (2–)3–5.5(–6)
cm long; seed scales 50–72(–80?) in number, 10–13
Conservation
× 7–10 mm, dark brown.
IUCN: DD
Distribution
Tsuga mertensiana (Bong.) Carrière subsp. grandi­
Pacific Coast Region of North America: from Alaska
cona Farjon, Proc. Kon. Ned. Akad. Wetensch., C,
to N California, also Rocky Mountains in interior
Bot. 91 (1): 39. 1988. Type: USA: California, Mono
British Columbia and Idaho.
Co., Mammoth Lakes, Twin Lakes, B. Willard 54
TDWG codes: 70 ASK 71 BRC 73 IDA ORE WAS 76
(holotype RM).
CAL

Conservation Description
IUCN: LC Short shoots apparent. Leaves usually crowded
and ascending above shoots. Seed cones 3.5–8.1 cm
Tsuga mertensiana (Bong.) Carrière subsp. merten­ long, 1.9–3.3 cm wide; seed scales (40–52)(–60?) in
siana (A. Henry) C. K. Schneid. var. jeffreyi, in number, 12–18(–20) × 10–15 mm, light brown.
Silva-Tarouca, Uns. Freil.-Nadelhölzer: 294. 1913.
Tsuga pattoniana (Balf.) Sénécl. var. jeffreyi A. Hen-
Distribution
ry, in Elwes & Henry, Trees Gr. Brit. Ireland 2: 231.
1907; Tsuga ×jeffreyi (A. Henry) A. Henry, Proc. USA: California (Siskiyou Mts., Sierra Nevada),
Roy. Irish Acad. 34: 55. 1919; ×Hesperotsuga jeffreyi S Oregon?
(A. Henry) C. N. Page, Notes Roy. Bot. Gard. Edin- TDWG codes: 76 CAL
burgh 45: 389. 1989. Type: not designated.
Conservation
Description
IUCN: NT
Foliage sprays short, asurgent; short shoots not dis-
tinct. Leaves more or less pectinately arranged. Seed
cones as in var. mertensiana (?). Tsuga sieboldii Carrière, Traité Gén. Conif.: 186.
1855. Type: Japan: [“in Japonia”], P. F. von Siebold
Taxonomic notes s.n. [comm. 1842 ex herb. Zuccarini No. 265]
(lectotype M).
This taxon was described from material said to have
come from Vancouver Island; however, recently it
Etymology
has not been found there, nor on the mainland of
British Columbia or in Washington, USA (Robert This species was named after Philipp Franz von Sie-
Van Pelt, University of Seattle, pers. comm. August bold (1796–1866), who studied the flora of Japan
when stationed there in the employ of the Dutch ovate-oblong, 5–6 × 3 mm, light yellowish brown,
East India Company. transparent.

Vernacular names Distribution

Southern Japanese hemlock; Tsuga, Toga matsu Japan: S Honshu, Kyushu, Shikoku, Yakushima.
(Japanese) TDWG codes: 38 JAP-HN JAP-KY JAP-SH

Description Ecology
Trees to 25–30 m tall, d.b.h. to 1.5–2.5 m; trunk Tsuga sieboldii grows in hills and mountains at al- 1019
straight or curved, rarely forked. Bark becoming titudes between 500 m and 1500 m a.s.l. (from 500
rough and scaly, breaking into square plates, dark m to 950 m on Shikoku). It grows on various soils
grey. Branches spreading horizontally or assurgent; derived from granitic or volcanic rock. The climate
crown broad conical in young trees, with drooping is moist temperate, with annual precipitation be-
leader, broadly domed or irregular, open and flat tween 1000 mm and 2000 mm, the winters are rela-
topped in old trees. Branchlets lustrous light brown, tively mild. Tsuga sieboldii is usually associated with
becoming grey with age, barely grooved between ap- conifers such as Abies firma, Pseudotsuga japonica,
pressed, decurrent, orange-brown pulvini, glabrous Chamaecyparis obtusa, Cryptomeria japonica, Pinus
or minutely pubescent. Vegetative buds ovoid or densiflora, P. parviflora, and Sciadopitys verticillata,
ovoid-conical, 2–2.5 mm long, not resinous, dark or- and with broad-leaved trees, e.g. Stewartia monadel­
ange-brown. Leaves parted above shoot, of unequal pha, Distylium racemosum and Trochodendron aral­
length, with shortest leaves assurgent above shoot, ioides. It rarely grows in pure stands.
some recurved, pectinate below the shoot, (0.5–)
1–1.5(–2) cm long, 2–2.5 mm wide, usually widest
Conservation
near truncate-emarginate apex, linear or ligulate-
linear, flattened, longitudinally grooved on upper- IUCN: LC
side; stomata on the abaxial (under) side in two
whitish bands separated by a midrib; leaf colour lus-
Uses
trous light green on upperside. Pollen cones crowd-
ed, 3–5 mm long, yellow when ripe. Seed cones The uses of the wood of Southern Japanese hemlock
numerous on outer branchlets of entire crown, short are mainly construction, carpentry and furniture
pedunculate or sessile, ovoid when closed, more glo- making. It grows less abundantly than Northern Jap-
bose with opened scales, (1.6–)2–2.5(–3) cm long, anese hemlock and was therefore not used on such
1.4–2(–2.5) cm wide, green or purplish green, rip- a large scale as a source for paper pulpwood. As an
ening to lustrous (pale) brown. Seed scales nearly ornamental tree it is grown in Japanese gardens and
circular, slightly convex, 8–10 × 8–12 mm at mid- parks, temple grounds and also in large pots. This
cone; abaxial surface striated or wrinkled, with im- species was introduced to Europe (the Netherlands)
prints of overlapping scales, glabrous; upper margin in 1850 by Von Siebold and is still in cultivation on
rounded or truncate, entire, undulate or (slightly) both sides of the North Atlantic, but uncommon. It
incurved; base auriculate-pedicellate. Bracts obtri- is slow growing and often makes a shrubby, spread-
angular, with denticulate upper margin, 2–3 mm ing tree with a dense crown.
long. Seeds ovoid, 2–3 × 1.5 mm, brown; seed wings
 Widdringtonia Endl., Gen. Pl. Suppl. 2: 25. 1842. Type: Widdringtonia nodiflora (L.)
Powrie [Brunia nodiflora L., lectotype by Powrie, 1972] (Cupressaceae).

Pachylepis Brongn., Ann. Sci. Nat. (Paris) 30: 189.

Distribution
1833, non Less. (1832). Type: Pachylepis cupressoides
(L.) Brongn. [Thuja cupressoides L.]; Parolinia Endl., Southern Africa: from the Cape to Malawi.
Gen. Pl. Suppl. 1: 1372. 1841, non Webb (1840). Type:
Thuja cupressoides L. [= Widdringtonia nodiflora
Key to the species of Widdringtonia
(L.) Powrie].
1020 1a. Seed cone scales strongly verrucose along mar-
Named after Samuel Edward Widdrington (1787– gins only, rugose on abaxial surface, with a
1856, formerly Cook), Commander of the Royal thick, robust boss (bract apex) 3
Navy. 1b. Seed cone scales smooth, rugose or irregularly
verrucose, with a thin, flattened boss 2
2a. Trees with monopodial stem and thick, fibrous,
Description
soft bark, never sprouting from base. Endemic
Shrubs or trees to 50 m tall, evergreen, monoecious; to Mt. Mulanje, Malawi W. whytei
trunk monopodial or multistemmed, resprouting 2b. Trees monopodial or multi-stemmed, with
from base or not. Resin in leaves and seed cones. thin, dense and hard bark, often sprouting from
Bark becoming fibrous soft or hard and fissured, base. Widespread from the Cape to Malawi
sometimes tesselated, exfoliating in long or short W. nodiflora
strips. Branches assurgent, ascending or spreading; 3a. Seeds angular-ovoid, with rudimental wings
foliage branches spreading to more or less erect. or nearly wingless. Endemic to the Cedarberg
Leaves scale-like, decussate on smallest branchlets, Mountains in Cape Province W. cedarbergensis
becoming more or less spirally arranged (bijugate) 3b. Seeds ovoid-oblong, with two wings exceeding
on thicker, leading (whip) branchlets (by twisting of 3–4 mm beyond the seed apex. Endemic to the
internodes), on ultimate branchlets appressed, ovate Willowmore District in Western Cape Prov-
to rhombic, increasing in size and becoming long ince W. schwarzii
decurrent on leading branchlets, upper leaf margins
minutely denticulate, apices often reflexed on lead-
Widdringtonia cedarbergensis J. A. Marsh, Bothalia
ing, thicker branchlets; amphistomatic, scale leaves
9 (1): 125. 1966. Type: South Africa: Cape Province,
eglandular. Pollen cones terminal, solitary; micro-
Clanwilliam Distr., Sederberge, near Middelberg
sporophylls 4–8, decussate, bearing 3–5 abaxial pol-
West Peak, H. A. Lückhoff s.n. (holotype PRE). Fig.
len sacs. Seed cones lateral, usually clustered (to 50
355
mature cones), sometimes terminal and solitary on
short shoots, initially consisting of 4 decussate, wide
Etymology
spreading green bracts and numerous ovules, ma-
turing in two growing seasons to irregular-globose, The species epithet refers to the Cedarberg Mts. in
serotinous but eventually wide openening cones. Western Cape Province, where this species is en-
Bract-scale complexes 4 (rarely 6), valvate, in two demic.
pairs of slightly unequal size and shape, thick woody,
the two largest more or less truncate, the smaller pair
Vernacular names
more acute, outer surface smooth to rugose, not or
variably verrucose, with an upcurved or recurved Clanwilliam Cypress, Clanwilliam Cedar, Cape Ce-
boss enclosing or exposing the bract apex, inner sur- dar; Clanwilliamseder, Sederboom (Afrikaans)
face with whitish seed scars at base. Columella short,
thick, angular, sometimes double. Seeds usually nu-
Description
merous, flattened or angular, with or without 2 wings
of unequal size. Seedlings with 2–4 cotyledons. Small trees to 20–22 m tall, often only 5–7 m; trunk
usually monopodial, stunted on exposed sites, to
4 species.
70 cm d.b.h. Bark on trunks becoming tesselated,
Distribution
exfoliating in small flakes. Branches spreading, con-
torted, forming a pyramidal crown in young trees, South Africa: Western Cape Province, Clanwilliam
spreading and irregular in older trees. Ultimate foli- District, Cedarberg Mts., in several localities re-
age branchlets 5–20 mm long, more or less quad- stricted to this small mountain range.
rangular in cross section, 1.5 mm wide, persistent. TDWG codes: 27 CPP-WC
Leaves on ultimate branchlets appressed, triangular-
ovate to rhombic, ca. 1.5 × 1 mm, increasing in size
Ecology
on older branchlets to 15 × 4 mm, keeled at base;
upper margins minutely denticulate; apex obtuse Remnant populations of this species are restricted
to acute; stomata on both sides in two marginal to rocky ridges and cliffs (‘krantzes’ or ‘kranse’) of 1021
lines; leaf leaf colour dull light green. Pollen cones Table Mountain Sandstone, avoiding other forma-
oblong, 3–5 × 1.5–2 mm; microsporophylls peltate, tions below and above it. The altitudinal range is
coriaceous, broadly triangular to ovate, bearing 3–4 (915–)1000–1500(–1650) m a.s.l. The trees are often
abaxial pollen sacs near base. Seed cones clustered protected by large boulders from frequent brush
on foliage branchlets, on reduced dwarf shoot axil- fires in the surrounding (secondary) fynbos vegeta-
lary to more or less spirally arranged leaves, some- tion. The annual rainfall is between 500–1000 mm
times terminal on short shoots, maturing in two and occurs mostly in the cold winter months, while
growing seasons to irregular-globose, 20–30 mm summers are dry and hot.
diam., serotinous but eventually wide openening
cones. Bract-scale complexes 4, thick woody, ob-
Conservation
long, max. 25 × 15 × 10 mm, curved slightly inward;
outer surface rugose, strongly verrucose along mar- There is a substantial literature concerned with
gins, with a prominent, 4–6 mm long, subapical, re- the conservation aspects of this relict species (for
curved boss enclosing bract apex, purplish brown or a selection, see Schellevis & Schouten in Farjon &
blackish brown; inner surface with angular, 4–6 mm Page, 1999: 91). Reports of a once existing forest of
long seed scars at base, yellow and lustrous brown. Clanwilliam cedars on the Cederberg are, by the na-
Columella short, thick, angular, to 6 mm tall. Seeds ture of European colonisation and the lack of earlier
(8–)10–20(–23) per cone, angular-ovoid, to 10 × 6 as well as contemporaneous written records, anec-
mm, trigonous, slightly flattened, lustrous yellow- dotal. Yet it is very likely that a historical reduction
ish brown ripening blackish brown, with dull lighter of more than 75% and perhaps as much as 95% has
hilum at base; wings 2, rudimental, forming thin, to occurred, resulting in a destroyed forest cover now
1 mm wide strips above widest part of seed, joining replaced by fynbos, a highly flammable vegetation
at seed apex. formation. The frequent fires prevent regeneration;
surviving trees are restricted to rocky ridges, ledges,
and outcrops mostly beyond reach of the flames.
Taxonomic notes
Attempts at replanting are continually being made
The name Widdringtonia juniperoides (L.) Endl., with low rates of success (Mustart et al., 1995) and
based on Cupressus juniperoides L., has been applied extensive ex situ conservation efforts are underway
to this species (e.g. Dallimore & Jackson’s A Hand­ in South Africa, backed by smaller scale plantings
book of Coniferae and Ginkgoaceae, ed. 4, 1966: 651). elsewhere.
However, the true identity of that name is enigmatic IUCN: EN (A1cd)
because the original description is too vague, the or-
igin of the plant unknown and no specimen or illus-
Uses
tration exist that could help identification and serve
as type. A new name was necessary, published in Clanwilliam cedar has been extensively logged for
the same year 1966 (op. cit.). The basionym Cupres­ its dense, easily workable and durable timber by
sus ­juniperoides L., together with all combinations European settlers. It was used for fence posts, con-
based on it, were listed with the incertae cedis in the struction of farm houses and sheds and later for
recent Monograph of Cupressaceae and Sciado­pitys carpentry, furniture, and cabinet making, until the
(Farjon, 2005a). resource dwindled and the last stands were almost
 destroyed by fires. In the nearby town of Clanwil- obtuse to acute, often reflexed on older branchlets
liam beautifully worked examples can be seen in and on branches with transitional leaves; stomata
the Anglican church (doors, pews, carved altar) and on both sides in two marginal lines; leaf colour dull
Courthouse. Today the few remaining trees are pro- light green. Pollen cones oblong, 3–5 × 1.5–2 mm;
tected from cutting for any purposes. Cultivation is microsporophylls peltate, coriaceous, broadly trian-
now undertaken primarily with a view of population gular to ovate, bearing 3–5 abaxial pollen sacs near
restoration and the species is consequently grown in base. Seed cones clustered (to 50 mature cones) on
local nurseries as well as in several botanic gardens. foliage branchlets, sometimes terminal on short
shoots, maturing in two growing seasons to irreg-
ular-globose, 10–20(–22) mm diam., serotinous
1022 Widdringtonia nodiflora (L.) Powrie, J. S. African but eventually wide openening cones. Bract-scale
Bot. 38 (4): 303. 1972. Brunia nodiflora L., Sp. Pl.: complexes 4, thick woody, oblong, max. 20 × 10 ×
199. 1753. Type: South Africa: Cape Province, leg. 7 mm, curved slightly inward; outer surface smooth
ign. s.n. [Clifford Herbarium, ‘Brunia 1’] (lectotype to rugose, not or variably verrucose, with a 1–4 mm
BM). Fig. 356 long, subapical, upcurved or recurved boss enclos-
ing or exposing the bract apex, (reddish) brown or
Thuja cupressoides L., Mant. Pl. 1: 125. 1767; Wid­ blackish brown; inner surface with angular, 3–4 mm
dringtonia cupressoides (L.) Endl., Gen. Pl. Suppl. 2: long, whitish seed scars at base, red-brown, black-
25. 1842. [Cat. Hort. Vindobon. 1: 209. 1842]. ish towards base. Columella short, thick, angular, to
Widdringtonia dracomontana Stapf, Bull. Misc. In- 5 mm tall, sometimes double. Seeds (3–)10–20(–30)
form., Kew 6: 206. 1918; Widdringtonia nodiflora (L.) per cone, ovoid, flattened, 5–6 × 3–4 mm without
Powrie var. dracomontana (Stapf) Silba, Phytologia wings, apically curved with a cuspidate tip, black-
68: 74. 1990. ish brown or black, with dull lighter hilum at base;
wings 2, of unequal size, up to 3 mm wide near seed
apex, not joining, brown, translucent.
Etymology
The species epithet means ‘with flowers at nodes’,
Distribution
but how Linnaeus could see this remains enigmatic.
Southern Africa: Malawi (Mt. Mulanje), W Mozam-
bique, South Africa, Zimbabwe, in mountainous ar-
Vernacular names
eas from S Malawi to the Cape.
Mountain Cypress, Cape Cypress; Bergsipres, Berg- TDWG codes: 26 MLW MOZ ZIM 27 CPP-EC CPP-
sapree (Afrikaans); Thaululo (Venda); Nwelelwen­ WC LES NAT OFS TVL-GA TVL-MP TVL-NP
taba (Zulu)
Ecology
Description
Predominantly in cool and wet mountain fynbos, of-
Shrubs or trees to 20–25 m tall; trunk monopodial ten in rocky outcrops and among boulders on sum-
or multistemmed, resprouting from base, to 50 cm mits, or in montane grassland often near streams,
d.b.h. Bark on trunks becoming fibrous but hard, to and in canyon woodland (‘kloofbos’), accompanied
10–15 mm thick, when fissured forming anastomo- by numerous fynbos genera (e.g. Leucadendron,
sing ridges, brown-grey, exfoliating in small strips. Leucospermum, Metrosideros, Protea, Restio) or
Branches assurging or ascending, forming a coni- Pteridium, Myrica pilulifera, and Poaceae, or form-
cal or pyramidal, eventually irregular crown. Ulti- ing pure stands. The altitude ranges from 100 m to
mate foliage branchlets 5–15 mm long, more or less 2590 m a.s.l. Soils are nutrient-poor, acidic, derived
quadrangular in cross section, 1 mm wide, persist- mostly from granite, quarzite or sandstone. The cli-
ent. Leaves on ultimate branchlets appressed, ovate mate varies from Mediterranean in the Cape region
to rhombic, 1.5–2 × 1–1.5 mm, larger and decurrent to subtropical with summer rains and tropical mon-
to ca. 10 × 4 mm on older branchlets, weakly keeled tane in Malawi.
at base; upper margins minutely denticulate; apex
 whip shoots; stomata on both sides in two marginal
Conservation
lines; leaf colour dull light green. Pollen cones ob-
This widespread species, which is capable of cop- long, 3–5 × 1.5–2 mm; microsporophylls peltate,
picing after above-ground destruction (fire), is not coriaceous, narrowly triangular to ovate, bearing
threatened with extinction. 3–4 abaxial pollen sacs near base. Seed cones clus-
IUCN: LC tered on foliage branchlets, sometimes terminal
on short shoots, maturing in two growing seasons
to irregular-globose, 20–30 mm diam., serotinous
Uses
but eventually wide openening cones. Bract-scale
No commercial uses have been recorded for this complexes 4, thick woody, oblong, max. 25 × 15 × 10
species. It may be readily coppiced and is probably mm, curved slightly inward; outer surface smooth 1023
used for firewood locally. Its only horticultural use or striate, strongly verrucose or tuberculate along
seems to be limited to plantings in botanic gardens; margins, with a prominent subapical, 5–8 mm long,
under glass where frost occurs and outdoors in re- thick, recurved boss enclosing bract apex, purplish
gions with a mild climate. It is suitable for planting brown or brown; inner surface with angular, 2–3 mm
in countries with a Mediterranean climate (and is long seed scars at base, yellow and lustrous brown.
planted in California) and should be used more of- Columella short, thick, angular, to 6 mm tall. Seeds
ten, provided it has first been assessed and found not 14–18(–20?) per cone, ovoid-oblong, 5–6 × 3–4 mm,
likely to become invasive. slightly flattened, curved towards cuspidate apex,
yellowish brown ripening blackish brown, with dull
lighter hilum at base; wings 2, on either side of seed,
Widdringtonia schwarzii (Marloth) Mast., J. Linn. exceeding 3–4 mm beyond retuse seed apex, un­
Soc., Bot. 37: 269. 1905. Callitris schwarzii Marloth, equal, thin, more or less translucent.
Bot. Jahrb. Syst. 36: 206. 1905. Type: South Africa:
Cape Province, Willowmore Distr., Kougaberge,
Distribution
R. Marloth 3614 (holotype SAM).
South Africa: Eastern Cape Province, Willowmore
District, Kougaberge, in canyons [‘kloofs’] draining
Etymology
into the Baviaanskloof and tributaries.
This species was named after “Herr E. Schwarz” who TDWG codes: 27 CPP-EC
drew Marloth’s attention to this tree.
Ecology
Vernacular names
Widdringtonia schwarzii occurs in rocky ravines on
Willowmore Cypress, Willowmore Cedar; Baviaans­ steep slopes or cliffs (‘krantzes’ or ‘kranse’) and in
kloofseder, Sapreehout (Afrikaans) (dry) river beds of canyons between (600–)900–
1200 m a.s.l. Trees can attain large size in rocky
streambeds of canyons, sheltered from ‘veld fires’,
Description
where they can form small groves. The climate is
Trees to 20–25 m tall, now rarely to 40 m; trunk mo- Mediterranean with mostly winter rain; the dry, hot
nopodial, to 1–1.5 m d.b.h. Bark on trunks fissured, summers are somewhat tempered by the sheltered
exfoliating in angular plates. Branches spreading, microclimate in deep, shady canyons, where deeper
contorted, forming a pyramidal crown in young ground water remains available under stream beds.
trees, more irregular in older trees. Ultimate foliage
branchlets 5–10 mm long, more or less quadrangu-
Conservation
lar in cross section, 1 mm wide, persistent. Leaves
on ultimate branchlets appressed, triangular-ovate In the past, settlers have exploited this species, which
to rhombic, ca. 1.5 × 1 mm, increasing in size on is capable of growing to large trees, to near extinc-
older branchlets to 10 × 4 mm, keeled at base; up- tion in all accessible areas. In recent years, new lo-
per margins minutely denticulate; apex acute, in- calities have been found in remote upper parts of
curved, often reflexed on thicker branchlets and on canyons in the Kouga Mountains, some of which
 harbour large trees. There is now much awareness colour dull light green. Pollen cones oblong, 3–6
of its conservation value and an active management × 1.5–2 mm; microsporophylls peltate, coriaceous,
programme (largely to prevent wildfires) is being broadly triangular to ovate, bearing 3–5 abaxial pol-
implemented. Further cutting of trees is prohibited. len sacs near base. Seed cones solitary or grouped
IUCN: VU (A1cd) (not clustered) on foliage branchlets, sometimes
terminal on short shoots, maturing in two growing
seasons to irregular-subglobose, 15–22 mm diam.,
Uses
serotinous but eventually wide openening cones.
Willowmore Cedar was used in the past for con- Bract-scale complexes 4 (rarely 6), thick woody, ob-
struction timber of local farmsteads and for furni- long, max. 25 × 15 × 8 mm, curved slightly inward;
1024 ture making. Depletion of the resource to a few trees outer surface smooth to rugose, not or variably ver-
in inaccessible locations terminated this use and the rucose, with a 1–4 mm long, subapical, upcurved or
remaining trees are now protected. In cultivation it recurved boss enclosing or exposing the bract apex,
is only present in a few botanic gardens and in some (reddish) brown or blackish brown; inner surface
private gardens in California and Oregon. It should with angular, 3–4 mm long, whitish seed scars at
be tried more often in summer-dry regions. base, red-brown, blackish towards base. Columella
short, thick, angular, to 5 mm tall, sometimes dou-
ble. Seeds 3–10(–18) per cone, ovoid, flattened, 5–7
Widdringtonia whytei Rendle, Trans. Linn. Soc. × 2–3 mm without wings, blackish brown or black,
London, Bot., ser. 2, 4: 60, t. 19. 1894. Widdring­ with dull lighter hilum at base; wings 2, of unequal
tonia nodiflora (L.) Powrie var. whytei (Rendle) size, up to 3 mm wide near seed apex, not joining,
Silba, Phytologia 68: 74. 1990. Type: Malawi: Mu- brown, translucent.
lanje District, Mulanje Highlands, Mt. Mulanje,
A. Whyte s.n. (holotype BM). Fig. 357
Distribution
S Malawi (Mt. Mulanje).
Etymology
TDWG codes: 26 MLW
This species was named after Alexander Whyte, who
explored Mt. Mulanje, Malawi, in 1891.
Ecology
Widdringtonia whytei is an important to co-domi-
Vernacular names
nant species in the Afromontane forest on Mt. Mu-
Mulanje Cypress, Mulanje Cedar lanje, which also includes Podocarpus milanjianus,
Cassipourea malosana, Ekebergia capensis, Olea
capensis, Polyscias fulva, Rapanea melanophloeos,
Description
and Xymalos monospora, and in the more fire-prone
Trees to 40–50 m tall; trunk monopodial, usually ecotone (with ericaceous scrub) to grassland, the
straight, to 1–1.5(–2?) m d.b.h. Bark on trunks be- closely related species Widdringtonia nodiflora. It
coming soft fibrous, to 2–4 cm thick, fissured, brown- is a successional species after fire (“periodic fire cli-
grey, exfoliating in long strips. Branches spreading max”), but unlike its congener, it does not coppice
or ascending, forming a pyramidal, eventually irreg- from (fire-caused) stumps and has to regenerate
ular or flat-topped crown. Ultimate foliage branch- from seed (Pauw & Linder, 1997). Thickets of Erica
lets (3–)7–20 mm long, more or less quadrangular in benguelensis which develop after fire offer protec-
cross section, 1–1.5 mm wide, persistent. Leaves on tion for cedar seedlings, leading to Widdringtonia
ultimate sterile branchlets appressed or with some whytei becoming the dominant tree until invading
free apices, ovate to rhombic, 1.5–3.5 × 1–1.5 mm, in- angiosperms succeed; however these have been pre-
creasing in size on leading branchlets (whip shoots) vented from doing so by the next fire at a cycle of
to ca. 10 × 4 mm, weakly keeled at base; upper mar- 100–200 years. Mt. Mulanje is a granitic batholith
gins minutely denticulate; apex obtuse to acute, of- rising through surrounding older sediments. The
ten reflexed on older branchlets and on whip shoots; soils are therefore largely rocky, acidic and shallow
stomata on both sides in two marginal lines; leaf except in colluvial pockets in gorges and valleys. The
altitudinal range is 1830–2550 m a.s.l. The climate is ings are to successfully contribute to the restoration
cool tropical montane, with abundant precipitation, of W. whytei.
much of it as fog. IUCN: EN (A1a-d, B1+2a-e)

Conservation Uses
This species is acutely threatened with extinction The wood of this species is highly valued for con-
because of excessive felling in the past 100+ years struction and building of houses as it is the only
and increased fire frequency (Chapman, 1995). Cer- sizable tree in the family Cupressaceae that occurs
tainly at present only moribund or severely exploit- naturally in a large surrounding region. Its wood, as
ed stands remain, mostly on less accessible sites. The of other members of the family, is decay and insect 1025
species is now officially protected on Mt. Mulanje, resistant. Large timber yielding trees are becom-
but enforcement proves to be difficult as the timber ing increasingly rare due to overexploitation of this
fetches high prices (Chapman, 1995; Pauw & Linder, valuable resource for more than a century (Chap-
1997). Population increase in villages surrounding man, 1995). Attempts at plantation forestry using
the mountain massif indirectly affects the remaining this species have been made, but have been much
populations through increased grazing of livestock less sucessful than the plantations of the exotic spe-
and the incidence of fires. Illegal timber extraction cies Cupressus lusitanica which grows faster. It also
continues to be a problem. Another potential threat appears that most of these plantations involve a
is the invasive pine Pinus patula, which is now being mixture of two species (Pauw & Linder, 1997), often
found in both grassland and forest on the mountain. with W. whytei at a disadvantage as it grows slower.
A more proactive strategy, involving the protection The wood of W. whytei is yellowish brown and of
of natural regeneration and perhaps planting, is ur- excellent quality for construction, general carpentry
gently needed to save this species from extinction. and joinery, wood panelling, flooring, and furniture
Ecologically it is at a disadvantage compared with its making. In colonial times some of its wood found
more adaptable congener on Mt. Mulanje, W. nodi­ its way to English interiors, nowadays there is no ex-
flora (Pauw & Linder, 1997). In past plantings, no port of this timber. This species does not appear to
distinction seems to have been made between the be in horticultural use, although it is present in a few
two species and this has to change if future plant- botanic gardens in South Africa.
 Wollemia W. G. Jones et al., Telopea 6 (2–3): 173. 1995. Type: Wollemia nobilis
W. G. Jones et al. (Araucariaceae).

Named after the Wollemi National Park (an Aborig- to 2–8 cm long, 2–5 mm wide. Adult upper canopy
inal name for the numerous canyons in the region). plagiotropic shoots initially near-vertical, becom-
ing horizontal and later pendulous, leaves opposite
or subopposite tetrastichous and twisted to present
Description
adaxial surfaces uppermost, narrowly oblong, coria-
See the species description. ceous, rounded, dull light to mid green, unequally
1026 amphistomatic, broad-based and decurrent; repeti-
tive growth units commencing with short scale-like
Distribution
leaves to 3 mm long, leaves increasing to 1–4 cm
As for the species. long, 4–8 mm wide. Pollen cones terminal on first-
order leafy plagiotropic shoots, to 11 cm long, 19 mm
diam., subtended by ca. 8 helically arranged broadly
Wollemia nobilis W. G. Jones et al., Telopea 6 (2–3): triangular to semicircular bracts 3–5 mm long, 3–5
174. 1995. Type: Australia: New South Wales, Wol- mm wide. Microsporophylls helically arranged,
lemi N. P., [exact locality kept secret], W. G. Jones dark red-brown, peltate, with a raised angular ter-
NSW 362731 (holotype NSW). Fig. 358, 359, 360 mination, with 4–9 elongate, pendulous pollen sacs.
Seed cones terminal on first-order leafy plagiotropic
shoots, usually borne on ascending branches above
Etymology
pollen cones, globular to broadly ellipsoidal, mid
The species epithet nobilis is Latin for noble and green, becoming brown and shedding individual
communicates the impression the tree gave its de- bract-scales at maturity, to 12.5 cm long, 10 cm diam.
scribers, but also refers obliquely to the discoverer Bract-scales flattened, laterally winged, 12–17 mm
of the first population of the species, David Noble. long, 14–22 mm wide, 3–5 mm thick, with a narrowly
triangular apical extension 6–12 mm long, 2–4 mm
wide at base. Seed scales wholly fused with and in-
Vernacular names
distinguishable from bracts. Seeds circumferentially
Wollemi pine winged, pale brown, 7–11 mm long, 5–7 mm wide,
6–9 mm wide including wing, remaining attached to
the scale. Germination epigeal, cotyledons 2.
Description
Evergreen monoecious trees to 40 m tall, frequently
Taxonomic notes
coppicing from base; trunks to 1.2 m diam, crown
slender, columnar, broadest at about a third of to- The discovery in 1994 of this unknown conifer and
tal height. Bark peeling in thin, fragile, equi dimen- the realization that it belonged to the family Arau­
sional dark red-brown scales on younger stems, cariaceae, but did not fit with either of the two
becoming densely covered with soft and spongy genera known in that family, caused considerable
darkening nodules or tubercules to 10 mm in diam. excitement among botanists. Further research indi-
Orthotropic shoots with helically arranged, decur- cated a slightly closer relationship with Agathis than
rent, narrowly triangular leaves 3–10 mm long, 2–4 with Araucaria (mainly based on molecular data),
mm wide at base. Juvenile and lower canopy plagio­ but many characters are distinct from either or simi-
tropic shoots horizontal, leaves opposite or sub­ lar to one but not the other in a combination unique
opposite, distichous and twisted to present adaxial to this new genus Wollemia. It soon became appar-
surfaces uppermost, linear to narrowly triangular, ent that a type of fossil pollen, Dilwynites, which had
chartaceous, rounded or obtuse, hypostomatic, deep long been known to petroleum geologists in Aus-
green above, glaucous below, broad-based and de- tralia, belonged to Wollemia and later some macro-
current; repetitive growth units commencing with fossils were ascribed to it as well, taking the fossil
short scale leaves to 3 mm long, leaves increasing record back to the Late Cretaceous (Turonian) as far
as is presently known (Kunzmann, 2007). The fossil searches have not yielded any other, more distant
record for Araucaria is substantially older (middle stands of this tree, so it is likely that the total popu-
Jurassic) and extinct representatives of the fam- lation does not exceed 100 mature trees and is prob-
ily Araucariaceae were present in the Triassic. The ably smaller. Counting individuals is difficult as the
chance survival of Wollemia in a gorge surrounded trees readily coppice from base or root meristems
by an environment hostile to it must count as one of and several trunks may belong to a single individual.
the remarkable events in the history of life. Access is difficult and the site is still kept a secret,
although by now (2010) there must be people who
have unauthorized knowledge of it. The species oc-
Distribution
curs entirely within the undeveloped Wollemi Wil-
Australia: New South Wales (Wollemi National derness of Wollemi National Park. Regeneration 1027
Park). occurs both from seeds and vegetatively and under
TDWG codes: 50 NSW-NS entirely natural conditions the species seems able to
survive. However, effects of human disturbance on
site and, in the longer term, possible effects of cli-
Ecology
mate change (e.g. drying of the permanent canyon
Emergent trees above warm temperate rainforest bottom creek) make these small populations, the
dominated by Ceratopetalum apetalum and Dory­ last to exist in a long declining trend as evidenced by
hora sassafras, in a deep sheltered gorge surrounded the fossil record, critically endangered.
by sandstone cliffs of the Triassic Narrabeen Group. IUCN: CR (D)
Tall eucalypt woodland dominated by Eucalyptus
piperita is adjacent on canyon walls and steep slopes,
Uses
giving way to dry open woodland up-slope. The soil
is sandstone-derived boulder alluvium, with high An extensive propagation and cultivation pro-
organic matter, some shale component, and a sub- gramme as well as marketing and PR have been
stantial basalt wash from higher reaches of small launched shortly after the discovery of this species.
tributary canyons. The local microclimate is wet, It has been available for sale since 2005 as a ‘living
with a permanent creek and an understorey domi- fossil’ in other words a curiosity; however, it appears
nated by ferns. to be suitable for large gardens in milder climates
(tolerating occasional light frosts, perhaps to zone 8
in Europe) and is an easy grower readily taking root
Conservation
from cuttings or scions. It is now well established in
This species was discovered in 1994 and since then horticulture and available from nurseries in many
three subpopulations, all within 2–3 km distance in countries.
a single canyon system, have been found. Extensive
 Xanthocyparis Farjon & Hiep, Novon 12 (2): 179. 2002. Type: Xanthocyparis
vietnamensis Farjon & Hiep (Cupressaceae).

Callitropsis Oerst., Vidensk. Meddel. Naturhist.

Taxonomic notes
Foren. Kjøbenhavn, ser. 2, 6: 32. 1864 (nom. ut. rej.,
Art. 56). Type: Callitropsis nootkatensis (D. Don) Since the publication of this genus and the two spe-
D. P. Little [Cupressus nootkatensis D. Don]. cies assigned to it (see below), more work has been
done to evaluate this new taxonomy. Notable in this
Greek: xanthos = golden or yellow; cyparis = cypress. context are the molecular (DNA) analyses by Little et
1028 al. (2004) and Little (2006). The first of these cladis-
tic analyses resulted in support for the unison of the
Description
two species, as proposed by Farjon et al. (2002), in a
Evergreen trees, monoecious, with fibrous bark ex- separate genus, related to Cupressus, but not closely
foliating in longitudinal strips; heart wood yellow- related to Chamaecyparis. However, it was discov-
ish, slow growing. Resin in leaves. Foliage branches ered that under the rules of the Botanical Code, an
spreading in plagiotropic sprays or drooping, forming earlier name for the genus had been proposed and
a pyramidal, conical or irregular flat-topped crown. should have been taken up: Callitropsis Oerst. This
Foliage of three types: juvenile linear leaves, transi- obscure and ambiguous name, which later had been
tional leaves and adult scale leaves. Juvenile leaves independently given to and often used for a conifer
present on seedlings only or also in mature trees, in New Caledonia, was subsequently proposed for
radially disposed in alternating whorls of four; foli- rejection under the Code, which has been effected,
age branchlets with this type of leaves always sterile. thereby reinstating Xanthocyparis as the legitimate
Transitional leaves present on seedlings only or also name for the genus. Meanwhile, Little (2006) had
in mature trees, decussate, divided in facials and lat- published a second analysis, this time with a wider
erals of nearly equal size; laterals weakly disposed in sampling of species. Adopting cladistic principles to
a plane; foliage branchlets with this type usually ster- use these new results in a classification, and appar-
ile. Adult leaves present in mature trees, decussate, ently unaware of the formal proposal to reject the
dimorphic in shape and size with laterals strongly name for future use, Little now proceeded to rename
flattened and disposed in a plane; foliage branchlets all American species of Cupressus to also be included
with this type often fertile. Leaves in whorls of four or in Callitropsis Oerst. This sollicited a response (Far-
decussate. Pollen cones terminal and solitary on lat- jon, 2007) rejecting this classification as a failure to
eral branchlets with small scale leaves, small; micro- consider the evolution of Juniperus (or other taxa)
sporophylls 10–16, decussate, peltate, bearing 2 (or 3) from a Cupressus(–like) ancestor. Under the (arbi-
relatively large pollen sacs. Seed cones terminal and trary) rule of cladistic classification, extant taxa can-
solitary on lateral branchlets with unmodified scale not have an ancestor-descendant relationship, they
leaves, opening wide. Bract-scale complexes 4 (or 6) are always ‘sister’ taxa.
in decussate pairs, fused at base, with connate upper
pair(s), spreading wide to release the seeds, valvate to
Key to the species of Xanthocyparis
subpeltate, with a prominent central umbo. Central
columella present or absent, small. Ovules axillary 1a. Juvenile, linear leaves and adult, dimorphic
to bracts, 1 to 5 per bract; seeds usually fewer, con- scale leaves commonly on the same foliage
centrated on upper pair (if 2 pairs) or middle pair branches of mature trees; facial scale leaves
(if 3 pairs) of scales, flattened, with two thin lateral ­eglandular or with an inconspicuous gland;
wings. Seedlings with 2 cotyledons. apex pungent X. vietnamensis
1b. Only adult, dimorphic scale leaves on foliage
2 species.
branches of young or mature trees; facial leaves
with a conspicuous gland; apex acute
Distribution
X. nootkatensis
Pacific Coast Region of NW North America; one lo-
cality in N Vietnam.
Xanthocyparis nootkatensis (D. Don) Farjon & or slightly reddish pollen sacs on lower margin. Seed
Harder, Novon 12 (2): 188. 2002. Cupressus noot­ cones maturing in 12–18 months, late caducous,
katensis D. Don, in Lambert, Descr. Pinus 2: 18. subglobose, 7–10(–13) mm with closed scales, from
1824; Chamaecyparis nootkatensis (D. Don) Spach, glaucous green ripening to purplish brown. Bract-
Hist. Nat. Vég. Phan. 11: 333. 1841; Callitropsis noot­ scale complexes 4(–6), decussate; upper pair usually
katensis (D. Don) D. P. Little, Syst. Bot. 31 (3): 474. larger than lower pair, parting and spreading wide
2006, (nom. ut. rej., Art. 56). Type: Canada: Brit- when mature, subpeltate, thick, oblong to weakly
ish Columbia, Vancouver Island, Nootka Sound, pentagonal in outline, joining at base, up to 9 × 8
A. Menzies s.n. (holotype not located, isotypes K, mm; abaxial surface smooth to rugose, slightly de-
MO). Fig. 361 pressed, with a central large umbo; adaxial surface
grooved and striated, brown, with light seed marks 1029
near base of upper pair(s). Columella more or less
Etymology
triangular or flattened, 1–2 × 1 mm. Seeds 6–10(–12)
The species epithet indicates the locality from where per cone, flattened, ovoid, 3–4 2–2.5 mm, reddish
it was first described: Nootka Sound between Van- brown, with a whitish hilum at base, surrounded by
couver Island and the mainland. 2 lateral, nearly equal thin wings 2–3 mm wide.

Vernacular names Taxonomic notes

Alaska Cedar, Alaska Yellow-cedar, Yellow-cedar, This species has been united with a newly discoved
Nootka Cypress, Sitka Cypress, Yellow-cypress cupressaceous conifer species in N Vietnam in
the new genus Xanthocyparis; the close relation-
ship between the two species has been confirmed
Description
by research using DNA sequence data. The name
Trees to 40–45(–60) m tall; trunk monopodial, up Xanthocyparis has been conserved under the Inter-
to 4–4.5 m d.b.h., multistemmed trees from layering national Code of Botanical Nomenclature (ICBN)
frequent at high altitudes or on bog margins. Bark against the earlier and ambiguous name Callitropsis;
soon flaky, on large trees exfoliating in thin strips, the latter has thereby been outlawed and is no longer
reddish brown weathering grey-brown. Branches available for any taxon.
spreading or curved down, higher order branches
drooping or pendulous, but top shoot drooping;
Distribution
crown becoming broadly conical in large trees. Fo-
liage branches drooping or pendulous, plagiotropic Pacific Coast Region of NW North America: from
branchlets alternating, forming tapering planate Prince William Sound in Alaska to the Siskyou
sprays, turning red-brown with dying leaves; ul- Mountains in California near the border with
timate lateral branchlets partly deciduous after ­Oregon.
4–5 years. Leaves decussate, imbricate, decurrent, TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
scale-like, coriaceous, 1.5–3 × 1–1.5 mm on ultimate
branchlets, up to 15 × 3 mm on leading shoots, di-
Ecology
morphic, of similar size; facials on branchlets nar-
rowly rhombic to lanceolate, acuminate-acute, Occurring across a vast latitudinal range, this spe-
appressed, with a conspicous, depressed, oval, non- cies is associated with different conifers from N to
active abaxial gland; laterals connate proximally, S and also along its altitudinal gradient (1–2300 m
spreading at apex of facials, conduplicate, broadly a.s.l.). Common in Alaska are Picea sitchensis and, at
falcate to lanceolate, recurved and incurved at acute, higher altitudes, Pinus contorta and Tsuga mertensi­
free apex, less conspicuously glandular or eglandu- ana; in British Columbia Abies amabilis, P. contorta,
lar; margins entire; stomata inconspicuous; leaf col- P. monticola, Thuja plicata and Tsuga heterophylla
our lustrous light green, on underside of branchlets and again at higher altitudes T. mertensiana; in
sometimes glaucous green. Pollen cones subglobose the Cascades at lower altitudes Abies amabilis and
to ovoid-oblong, 3–5 2–2.5 mm; microsporophylls A. procera, at higher altitudes A. lasiocarpa, P. albi­
subcordate, with entire margins, with 2(–3) yellow caulis and T. mertensiana; and in its southernmost
 outlier the Siskiyou Mountains it is associated with ­ nder the new taxonomy and it has become ×Cupro­
u
A. magnifica, Calocedrus decurrens and Picea brew­ cyparis leylandii (A. B. Jackson & Dallimore) Farjon,
eriana. A shrub-layer, in which ericaceous species as minimal a change as possible (Farjon et al., 2002).
often dominate, is usually well developed. Character- This new name has now been adopted by the Royal
istic are stands, sometimes pure, or with P. contorta Horticultural Society’s (RHS) Conifer Register of
or T. mertensiana, forming a forest ecotone around recommended names for conifers in cultivation
bogs or near the tree line, with scrubby growth forms (Springate, 2009). The hybrid plants are sterile and
predominant on organic soils (derived from peat) in must be propagated vegetatively as clones; their hy-
both habitats. As with all conifers in these forests, brid vigour (heterosis) has earned notoriety as they
competition on better sites favours faster and taller have been widely used for hedges that subsequently
1030 growing species, which means X. nootkatensis is not grew out of proportion and often control. Garden
codominant there and gets eventually pushed out crosses with two other New World Cypresses (Cu­
to sites with shallow soils or nutrient deficient soils. pressus arizonica var. glabra, C. lusitanica) are also
However, it is also in much of its range probably the in cultivation, but much less common.
longest-lived conifer, with ages well over 1500 years
verified and in excess of 2000 years, and probably
even up to 3500 years (trees of this inferred age are Xanthocyparis vietnamensis Farjon & Hiep, Novon
found to be hollow). It could therefore well exhibit 12 (2): 180. 2002. Callitropsis vietnamensis (Farjon &
a strategy tied in with episodal disturbance events Hiep) D. P. Little, Amer. J. Bot. 91: 1879. 2004, (nom.
in which certain individuals outlive all competitors ut. rej., Art. 56); Cupressus vietnamensis (Farjon)
and only regenerate after this rare event. The cli- Silba, J. Int. Conifer Preserv. Soc. 12 (2): 100. 2005.
mate, especially in the very maritime near coastal Type: Vietnam: Ha Giang Prov., Quan Ba District,
strip and on ocean-facing slopes and islands, is cool Bat Dai Son Provincial Protected Area, D. K. Harder
and very wet. et al. 6091 (holotype OBS). Pl. 42, Fig. 362

Conservation Etymology
Although ‘dieback’ (with largely unknown causes) The species epithet denotes Vietnam, the country in
has been reported in parts of its range, this species is which it is endemic.
widespread and occurs in many places that are still
remote.
Vernacular names
IUCN: LC
Golden cypress; Bach vang (Vietnamese)
Uses
Description
The wood of this slow growing species is extremely
durable and valuable, being used for boat building Trees to 10–15 m tall; trunk monopodial, up to 50 cm
and other maritime building and generally for out- d.b.h. Bark on the trunk of larger trees becoming soft
door construction in cool and wet climate. Most of and fibrous, brown to grey-brown, exfoliating in nu-
the high quality logs are being exported and a large merous thin strips. Branches spreading more or less
proportion of the timber goes to Japan. In horti­ horizontally; forming a pyramidal crown in young
culture usually known as Nootka cypress, this spe- trees but a spreading, irregular or flat-topped crown
cies is often used as an ornamental and a number of in old trees. Foliage in crowns of both small and larg-
cultivars are known. It is also the maternal parent er trees of two or three types: predominantly with
of the intergeneric hybrid Xanthocyparis nootkaten­ adult leaves, also with juvenile leaves, often also with
sis (syn. Chamaecyparis nootkatensis) × Cupressus transitional leaves. Foliage sprays with adult leaves
macrocarpa or Leyland cypress which originated flattened; ultimate branchlets spreading at 30–45°,
in cultivation at Leighton Park in England in 1888. 5–20 mm long, 1.5–3 mm wide. Foliage sprays with
This has consequences for the botanical name of the juvenile leaves bushy, less branched; branchlets with
Leyland Cypress. The International Code of Botani- transitional leaves more like those with adult leaves.
cal Nomenclature (ICBN) requires it to be changed Adult leaves decussate, short decurrent, imbricate,
 1031

3
7
9
6

5
2

Pl ate 4 2 . Xanthocyparis vietnamensis. 1. Branch with adult foliage and seed cones. 2. Branch with juve-
nile and adult foliage. 3. Adult leaves. 4. Juvenile leaves. 5. Surfaces of juvenile leaf. 6. Pollen cone. 7. Micro-
sporophyll with pollen sacs. 8. Seed cones. 9. Seeds.
 dimorphic, on (sub)ultimate branchlets 1.5–3 × 1–1.3 angiosperms are species of Acer, Carpinus, Litho­
mm (the laterals slightly longer than the facials); fa- carpus, Quercus, and Ulmus; frequent are Pistacea
cials narrowly ovate-rhombic, keeled, more or less weinmannifolia and Platycarya strobilacea. In a sec-
appressed; laterals conduplicate, spreading free from ond stratum under the ca. 20 m tall canopy occur
the leaf above at ca. 30°, straight or falcate; margins frequently species of Elaeocarpus, Eriobotrya, Sor­
of both types minutely denticulate except towards bus, Schefflera, and many others. Shrubs and herbs
acute or pungent apex; stomata on adult leaves in- abound, among the latter are numerous species of
conspicuous, mostly adaxial; glands inconspicuous. Orchidaceae, terrestrial as well as epiphytic, some-
Transitional leaves similar to adult leaves but longer times determining the aspect of the ground cover
(5–7 mm), lanceolate; laterals spreading wider at 45°. vegetation (Averyanov et al., 2002). Ferns and es-
1032 Juvenile leaves in whorls of four, decurrent, with dis- pecially bryophytes are similarly abundant, both as
tal part spreading at nearly 90° and proximal decur- lithophytes and as epiphytes. The limestone ridges
rent part 4–5 mm long; distal free parts 15–20 × 1,5–2 on which Xanthocyparis occurs are extremely erod-
mm, linear; margins entire, tapering to a fine point; ed, composed of hard, marble-like rock outcrops
stomata in two whitish bands on abaxial side only. interspersed with thin soil pockets. The altitude of
Pollen cones 2.5–4 × 2–2.5 mm, oval-terete; micro- the ridges where this conifer grows is from 1050 m
sporophylls 10–12, peltate, ca. 1 × 1 mm, each bearing to 1330 m a.s.l. The climate is subtropical, but damp
abaxially 2(–3) yellow pollen sacs. Seed cones green, and wet much of the year (cloud forest).
turning dark or dull brown, subglobose, 9–11 ×
10–12 mm when open, some more or less persistent
Conservation
after seed dispersal. Bract-scale complexes 4(–6),
decussate, fused at base; upper and larger pair con- This species is restricted to a few localities in close
nate distally, then spreading, valvate to subpeltate; proximity, mostly in inaccessible sites on steep lime-
lower pair oblong; all widest distally, with rounded stone ridges. In some localities trees are cut, pre-
but irregular upper margin; outer surface rugose sumably for local use, in others no such disturbance
or radiately furrowed from a prominent, 1–2.5 mm could be found, possibly due to inaccessibility. The
long, subapical umbo; inner surface reddish brown total extent of occurrence (EOO) appears to be less
marked proximally with whitish seed scars; a small than 50 km2 within which the species occupies ridge
columella present or absent in the centre. Seeds up tops and summits only. Although difficult to assess
to 8–9 per cone, ovoid or irregular, flattened, 4.5–6 precisely due to the difficult terrain, the total area
× 4–5 mm including two lateral wings, light brown of occupancy (AOO) is estimated to be less than
or reddish brown, with whitish hilum at base; wings 10 km2 (Averyanov et al., 2002). Regeneration has
0.5–1 mm wide, thin menbranous, lighter coloured. been observed in habitat. An earlier estimate (Farjon
et al., 2002) put this species at Critically Endangered
(CR), as did Averyanov et al. (2002), but objective
Distribution
evidence of serious decline has not been given.
Vietnam: N Vietnam, Ha Giang Province, very lo- IUCN: EN (B2 a-b)
calized in the Bat Dai Son mountain system near the
Chinese border.
Uses
TDWG codes: 41 VIE
The Golden cypress produces fine, yellowish brown,
very hard, fragrant timber. The majority of trees
Ecology
seen on the ridges are small or contorted and would
This species occurs in mixed angiosperm-conifer not be suitable for timber of any trade value other
forest with the conifers Amentotaxus argotaenia, than local use. To date, no direct information of its
Nageia wallichiana, Pseudotsuga sinensis (domi- actual or potential uses is available. Cultivation is
nant), Podocarpus brevifolius, and Taxus chinensis being undertaken locally with an objective of ex situ
and numerous, mostly small leaved angiosperms. conservation; a few botanic gardens in the UK and
The conifers appear to be restricted to the ridges USA also grow it for this reason.
and summits of the mountains. Dominant among
GLOSSARY

abaxial In a position removed from an axis or rounded by a pericarp formed by the carpels;
stem (also dorsal). the seeds are consequently enclosed in an ovary
acicular Needle-shaped; long, narrow, straight or which develops into a fruit of some kind (com-
slightly curved, slender, and not or only slightly pare gymnosperms). 1033
flattened. anthesis (in gymnosperms) The action or period
acroscopic Facing or directed towards the apex of pollen dispersal and pollen reception (‘flow-
(here of the foliage branch). ering’).
acuminate Tapering gradually to a well defined, AOO (area of occupancy) Term used in IUCN
sharp point. Red List criteria to describe the estimated area
acute Sharply pointed. (in hectares or square kilometres) actually cov-
adaxial In a position nearest an axis or stem. ered or occupied by a species. Compare EOO.
adnate Joined with another organ of a different apex In plant morphology, the terminal point of
kind; e.g. the seed wing with a seed (compare a growing structure like a shoot or a leaf. The
connate). adjective is apical; a conifer cone that is apical
adult leaves Here the type of green leaves that ap- grows at the apex of a shoot (but very few are).
pears and is predominant on primary (or old) apiculate Ending abruptly in a small, distinct
branches of mature trees, usually in the crown point.
and fully exposed to sunlight (compare juve- apiculus A small, abrupt and/or distinct point at
nile leaves). the apex.
afforestation The planting or seeding of trees in apophysis The well defined apical part of ovulif-
an attempt to create ‘forest’. What is meant with erous scales in the seed cone marking the first
the term forest has changed over time; in this stage of growth in Pinus; it is rarely differenti-
context it now usually refers to plantations used ated in the other genera of Pinaceae.
as a source of timber. appressed Pressed close to or lying flat against an-
aggregate Grouped closely or densely together, other organ (e.g. the shoot or other scales).
clustered. arboretum (plural: arboreta) A park or garden
allopatric The occurrence of populations or spe- dedicated to a living collection of planted trees
cies in geographically separated areas. and shrubs.
amenity planting Planting of ‘ornamentals’ and archegonia (singular archegonium) In gymno-
trees for general use in public spaces, e.g. street sperms, the unfertilized female ‘sex organs’, i.e.
trees, flower borders, and shrubs in parks, high- the parts of the ovule that contain the female
way intersection plantings, etc. nuclei with a haploid number of chromosomes.
amphistomatic Stomata on both abaxial and Gymnosperms have several archegonia per
adaxial sides, on equifacial leaves on all three ovule.
or four faces; equally distributed, or more often aril (Latin arillus) A fleshy or succulent (cup-like)
with two primary bands on two adjacent faces structure that completely or partially envelops
(compare epistomatic and hypostomatic). a seed and is derived from the integument
ampulliform Shaped like a flask (Latin ampulla), (compare epimatium and drupe).
with a wide body and a narrowing neck below ascending Rising obliquely or curving upward.
the apex. a.s.l. Above sea level.
anastomosing Conneting so as to form a network asperous Harsh to touch or handle; here due to ri-
(compare reticulate). gidity of pungent leaves.
angiosperms Flowering seed plants with ovules association (in vegetation science) A fundamental
attached to carpels, developing into seeds sur- unit of vegetation, which can be distinguished
 from other such units on the grounds of its flo- bole The straight trunk of a tree.
ristic composition. boreal (Greek Boreas = god of the north wind) Of
assurgent Bent near the base, then steeply rising northern regions of the world.
to a (nearly) vertical position. bract A leaf-like or scale-like appendage, usually
auriculate (auricled) With ear-like lobes at the subtending a fertile structure such as a seed
base or lower sides. scale. In some conifers, the seed scale is fused
axil The angle between a bract, branch, or leaf and with the bract, rudimentary, or altogether ab-
the axis from which it arises (i.e. rachis, shoot, sent, by which the ovules (seeds) are situated
branch, or stem). in the axil of bract and cone axis, or sometimes
axillary Positioned in the axil. solitary on the apex of the axis.
1034 basionym The name-bringing or epithet-bringing bract-scale complex Term used in Cupressaceae
synonym of a newly proposed combination. for the scales of seed cones; these are techni-
bifacial With two distinct, opposite faces. If flat- cally bracts, but transformed by intercalary
tened in that way, the organ, e.g. a leaf, has its growth to form the cone scales and enclose the
margins bordering these faces (see also dorsi­ seeds. In some complexes there is a rudimenta-
ventral and compare bilateral). ry seed scale present (in related Sciadopityaceae
bifid Deeply cleft at apex, so as to form two sharp it is fully developed), in most the true seed scale
teeth. does not develop.
bijugate An arrangement (of leaves) in which buttressed Shaped like a buttress; here the broad-
there is a doubling of the numbers 2 + 3 in ened base of a tree trunk, usually continued
normal (unijugate) spiral arrangements; the bi­ downward in the above-ground parts of roots.
jugate arrangement of 4 + 6 ‘parastichies’ is in caducous Falling off at an early stage.
fact an opposite-decussate arrangement with a callous Being hardened and thickened.
‘twist’. cambium Primary tissue (see meristem) forming
bilateral A symmetry in which similar parts secondary tissue. There are two cambia: one in
(sides) are arranged on opposite sides of a me- the outer part of the stem (cork cambium) and
dian axis; in leaves: flattened with two equal one just outside the wood of the stem; it forms
sides (not faces; compare bifacial and dorsi­ secondary wood on its inner side and second-
ventral). ary bark on its outer side.
bilobate With two lobes; usually a terminal part canaliculate With a single, longitudinal groove.
divided in two. candelabra crown A tree crown conforming to
binomial A name of a thing consisting of two sep- Rauh’s model but with few first order branches
arate words; in biology it refers to the names of in pseudo-whorls restricted to the upper part of
species, which are always the name of a genus the trunk and with few higher order branches
written with a capital first letter followed by an limited to the ends of these.
epithet written in lower case. canopy The upper crown level in a closed forest,
biodiversity The total number of taxa (or some- where the crowns of the tallest trees meet to
times genetic diversity) known to exist in a giv- form a more or less continuous cover.
en area; the greater this figure turns out to be, capitula (Latin = small head; plural: capitulae) A
the higher the region’s biodiversity. dense, compact cluster of flowers. In conifers,
biogeography The study or science of the geo- this refers to pollen cones, not flowers.
graphical distribution of organisms and its carinate Keeled; ridged lengthwise.
causes and history. cataphyll A reduced, scaly primary leaf; in pines
biome A major ecological community type, relat- they form winter buds, enclosing an entire
ed a to major climatic zone. shoot (with leaves and internodes not yet elon-
biota (Latinised form of Greek biotē = life) The gated) of the next growing season. On expan-
totality of life (plants, animals, fungi etc.) in a sion of the shoot each cataphyll subtends a
region. dwarf shoot with secondary leaves (needles).
bisaccate (in pollen) Having two sacci or air blad- caudate Elongated, resembling a tail, or tail-like
ders giving the corpus of the pollen buoyancy appendage.
in the air.
chaparral Term used in the southwest of the USA variables) of at least three different organisms
and Mexico to describe a dense vegetation of (representing species or higher ranks of taxa),
shrubs, some of which are evergreen, which showing the inferred phylogenetic relation-
grows in coastal regions with a dry summer ships of these characters based on cladistic
and a moist winter climate, and often forms a principles.
distinct vegetation belt (see also maquis). clavate Club-shaped, i.e. with a broader or thicker
character A feature of an organism which has var- distal part.
iable properties. climax A supposedly terminal stage in the succes-
character state The variable property of a charac- sion of vegetation types from pioneer coloniz-
ter. Leaf length is a character, the actual meas- ers to mature vegetation. In fact, all stages in
urement given in cm or mm is the character the succession are temporary since they are all 1035
state. subjected to natural disturbances, including the
chlorophyll A green pigment, contained in chloro­ terminal or climax stage.
plasts in leaf or sometimes stem tissue of plants, cline A graded series of morphological (or physi-
responsible for photosynthesis. ological) differences exhibited by a group of
chloroplast A microscopic body (organelle) in- related organisms, usually along a line of envi-
side a plant cell that contains chlorophyll. ronmental or geographic transition.
chromosome A usually linear body of chromatids, codominant Dominating together with other spe-
containing nucleoprotein and DNA, the mole- cies in a given vegetation; trees are codominant
cule responsible for the hereditary code, in the when they form part of the main canopy of a
cell nucleus; the number of chromosomes usu- forest.
ally remains constant in a species. columella (Latin = little column) In conifers a
ciliate Fringed with regularly arranged hairs on spiky protrusion in the centre (axis) of the in-
the margin; the hairs are usually fine and of side of a seed cone, especially prominent in
equal length. those of the genus Callitris (Cupressaceae).
CITES Convention on the International Trade in concave Hollowed or rounded inward like the in-
Endangered Species of Wild Fauna and Flora. side of a bowl (compare convex).
The signatories to this convention (1976) are conduplicate Folded together lengthwise.
nation states; they decide at regular conferences cone In conifers, referring to a structure bearing
of the parties to restrict or prohibit cross-­border either pollen sacs containing pollen, or ovules,
trade (= movement by people) of species listed which after fertilization may develop into seeds.
on Appendices I, II, or III (with I the most re- Conifer cones can be simple, consisting of an
strictive). axis with one type of scales, or compound, with
clade A section of a cladogram which indicates a two types of scales, only one of which bears the
monophyletic (holophyletic) group of ‘termi- ovules. Pollen cones in conifers are simple; seed
nal’ taxa (2–∞). cones can be compound or simple.
cladistics A formalised method of character com- congener A species in the same genus as another
parison developed by the German entomologist species.
Willi Hennig in 1950; it only considers shared conifers (cone-bearers) To this taxonomic group
derived characters to inform phylogeny and its belong those woody gymnospermous plants
aim is to construct a hypothesis of phylo­genetic (usually trees) of which the seed bearing scales
relationships between groups of organisms are united in a cone. The conifers (including
(usu­ally recognized taxa). forms with extremely reduced cones, such as
cladode A branch-like or shoot-like organ not be- Podocarpus, or those with no apparent cones,
ing a green leaf, but functioning as such (com- such as Taxus) are a monophyletic group of
pare phylloclade). In Sciadopitys true leaves plants derived from a single common ancestor.
form small scales in the axil of which arise nee- connate Joined with another organ of a similar
dle-like, seemingly paired, and fused cladodes, kind, e.g. a leaf with another leaf (compare ad-
functioning as leaves. nate).
cladogram A branching diagram resulting from convergence The evolution towards similar mor-
a cladistic analysis of characters (i.e. discrete phology not as a result of common descent, but
 of adaptation to similar environments or life d.b.h. Diameter at breast height; used to measure
strategies. the girth of a tree trunk above a buttressed base
convex Curved or rounded like the exterior of a or roots; for very large buttressed trees meas-
sphere or circle; here used for the shape of the urement should be taken at actually higher
cone scales as viewed from the abaxial side (than the usual ca. 1.3 m) levels.
(compare concave). deciduous Falling off or shed seasonally or at a
copal A kind of resin; commercially referring to certain stage of development in the life cycle.
resins of leguminous and araucarian plants that decumbent (Latin = lie down) Growing low over
produce hard varnishes. the ground or surface of a rock.
coppice Regrowth from the stump of a tree after decurrent Running down a stem from a leaf base.
1036 cutting (or burning), producing new stems and decurved Curved downward (compare recurved).
branches. The stump after cutting is called cop- decussate Occurring in alternating pairs; leaves
pice stool; in many species the stems will re- are decussate if one opposite pair is placed
grow even after repeated cutting. slightly higher than the adjacent opposite pair
cordate Heart-shaped (subcordate = somewhat and oriented ± 90° from it (compare ternate).
heart-shaped). deflected (Latin deflexus) Bent or turned abruptly
coriaceous (Latin corium = skin) Resembling downwards.
leather: leaves with a leathery, tough texture, dehiscent Referring to the manner of opening of a
mainly from thickened dermal layers. vessel containing seeds or spores; in conifers it
cortex Layer of` cells (parenchyma) in plant roots pertains to the pollen sacs or to some types of
and stems, situated between the central vascu- globose seed cone.
lar cylinder and the epidermis or surface layer. deltoid triangular with the sides more or less equal
costate Ribbed, having one or more primary veins and the apex uppermost.
or ridges. dendrochronology The science of dating events
cotyledons First, or embryo leaves, as found in the and variations in the environment in former
seed and which (usually) unfold after germina- periods by comparative study of growth rings
tion. in the woody stems of trees.
crenate (crenately lobed) Scalloped, lobed like the dendrological Of dendrology, the science or study
surface of a scallop. of trees.
cultivar A form of a plant (not a botanical vari- dentate Having the margin cut with sharp, salient
ety!) in cultivation, the characteristics of which teeth, directed outward or straight.
are perpetuated by vegetative propagation. denticulate Finely or minutely dentate.
cuneate Wedge-shaped; in the descriptions often dermal Of the skin (Greek derma = skin), in coni-
combinations are used, e.g. obovate-cuneate, to fers it can refer to bark or more commonly to
describe either a range of shapes or intermedi- leaf surfaces or outer layers (see epidermis).
ates. determinate Referring to growth of a structure
cupular Cup-like, similar in shape to the cupule which definitely ends with the completion of
of an acorn. that structure. Conifer cones represent shoots
cusp The usually well defined pointed apical ex- with determinate growth, while foliage shoots
tension of a leaf (but here also of the bracts of can and often will resume growth in the next
seed cones). growing season. In some abnormal cases, cones
cuspidate With a cusp, i.e. a (short or long) well are not determinate and a foliage shoot may
defined and tapering apical point. grow from their apex (compare proliferation).
cuticular Referring to a usually waxy surface layer dichotomous Like a system of branching in which
on the exterior of the epidermis, consisting of the main axis forks repeatedly into two branch-
a protective substance (cutine); it causes the es (dichotomy).
shine on conifer leaves. dilated (Latin dilatatus) Expanded, broadened, or
cyathiform Shaped like a beaker (in outline) (Lat- widened (sometimes also swollen).
in cyathus = beaker), i.e. a wide-mouthed and dimorphic Of two forms or shapes; an organ, e.g. a
projecting-lipped drinking vessel supported on leaf, has two different forms (2 character states)
a standard.
 occurring on the same plant or shoot (compare ecotone The transitional zone between two types
monomorphic and polymorphic). of vegetation or ecological communities, usu-
dioecious In gymnosperms this pertains to fe- ally with (competing) species of both types and
male and male reproductive organs, commonly often with species not occurring in either of the
cones, being present on different plants (see two types.
also monoecious). ecotype A plant form with morphological char-
diploid Cells containing two sets of chromosomes acters that are modified by the environment in
(2n = x). The nuclei of these cells contain sets which the individual plant grows; such char-
of paired chromosomes of equal form and size. acters are not inherited and therefore are not a
Such cells have a doubling of the base (haploid) good basis for taxonomic distinction.
number of chromosomes, the result of sexual emarginate Shallowly notched at the apex. 1037
fusion of female and male haploid nuclei. A embryo (in plants) The young sporophyte of a
rare further doubling of the diploid number seed plant, usually consisting of a very small
leads to tetraploidy (Greek tetra = four) and in plant with a primary bud (plumule), radicle,
one species (Sequoia sempervirens) hexaploids and cotyledons.
(Greek hexa = six) were found, which amounts emend. (Latin emendavit = changed by) Indicat-
to a tripling of the diploid number. ing an alteration of content or circumscription,
disjunct (of distribution) Occurring in areas followed by citation of the author(s) responsi-
widely separated from each other. ble for the change.
dispersal The spreading of organisms to areas oth- emergent In the context of trees referring to indi-
er than where they originated. viduals rising above the forest canopy.
distal Occurring nearest the apex or terminal part endemic (endemism) Referring to the occurrence
(compare proximal). of a taxon in a limited area ‒ for example, on
distichous Disposed in two vertical or upright a single island. Taxa limited to a single coun-
rows. try are also considered endemic, although the
divaricate (from Latin divaricare = ‘to straddle’ as term becomes fairly meaningless biologically
in riding a horse) Divergent branching, with for widespread species limited to countries that
two branches splitting from the apex of a single cover nearly half a continent.
branch or stem. endophyte An organism that grows within (in the
DNA (deoxyribonucleic acid) The macro-mol- context of trees a branch, root, or stem).
ecule in cells of organisms that contains the entire Without teething or division, with even
genetic information and acts as a template to margin.
build new cells and therby (multicellular) new EOO (extent of occurrence). Term used in IUCN
organisms (compare RNA). Red List criteria to estimate the total range (in
dolabriform Shaped like an axe. km2) within which a species is found to occur
dorsiventral The condition of possessing upper naturally (excluding introductions), usually by
and lower faces of an organ. If flattened in that connecting the outermost points of present oc-
way, the organ, e.g. a leaf, has its margins bor- currence. Within the EOO is the AOO, which
dering these faces (see also bifacial). is always smaller.
drupe A fleshy, one-seeded, indehiscent fruit with epicormic (Sprouting) from dormant buds on the
a stony endocarp (e.g. cherry, peach). It is de- trunk lying within the bark.
rived from the carpels of angiosperms (com- epidermis The true cellular outer skin of a plant
pare aril and epimatium). (leaf) below the cuticle, made up of a single
ecology The study or science of ecosystems, or of layer of cells.
the functional relationships of organisms with epigeal germination The hypocotyl of the seed-
their environment and with each other. ling elongates and lifts the cotyledons above
ecosystem The supposed totality of the functional the soil surface, where they unfold (compare
relationships of organisms with their environ- hypogeal).
ment and with each other living in a given area epimatium The outer layer of tissue that (partly)
and habitat. covers a seed in many species of the family
 ­ odocarpaceae; it is derived from the seed scale
P exserted Protruding; here usually pertaining to
(compare aril and drupe). the bracts of seed cones projecting beyond the
epiphyte A plant that grows upon another plant upper margins of the seed scales.
(usually a tree) using it for support only. Plants ex situ (Latin = outside [its] place) In the context
that are ‘accidentally’ growing on trees because of plant species conservation, any effort to con-
seed germinated there, but of which the usual serve individuals by growing them in localities
habitat is terrestrial, are not (obligate) epiphytes outside the natural range of the species is con-
in an ecological sense. No obligate coniferous sidered ex situ conservation (compare in situ).
epiphytes are known. exuding Issuing a substance, such as resin, through
episodal Occurring at irregular intervals in time a surface layer of tissue; the product is called
1038 (episodes). Here referring to stochastic distur- exudate.
bance events from natural causes that can reset facials (facial leaves) The paired, opposite scale
the successional phases of forest regeneration leaves on a plagiotropic foliage branchlet (esp.
in an area to a pioneer phase. in the family Cupressaceae) that face either up
epistomatic Stomata (almost) restricted to the or down, they are flanked by the differently
adaxial side of the leaves (the dorsal side mor- shaped laterals.
phologically); on plagiotropic lateral shoots falcate Sickle-shaped (falcate-linear = with two
the stomata are facing downward by orienta- nearly parallel sides).
tion of the leaf bases, petioles or pulvini (com- fastigiate A branching habit in trees in which the
pare hypo­stomatic). branches are directed upwards (ascending),
epitype (Latin epitypus) A specimen or illustration not spreading out from the trunk.
selected in addition to a holotype (or neotype farinose Covered with a flowery or meal-like
or lectotype) to serve as an interpretative type powder.
if the designated type material or other ‘original ferruginous With the reddish brown colour of
material’ cannot be identified for the purpose rusty iron.
of precise application of the name of a taxon. figured In sawn and planed or turned and pol-
equifacial With equal faces; i.e. the 3 or 4 faces of ished wood the decorative patterns and shapes
those leaves that are quadrangular or rhombic caused by growth rings and other structures in
in cross-section. relation to the plane of sawing or turning.
ericoid (as in Erica) Belonging to (or similar to) fimbriate Fringed with long hairs.
Ericaceae; also an adjective used to describe flabellate Fan-shaped (though in cone scales of
a vegetation dominated by such plants, or by conifers not entirely flat but more or less con-
those with a similar life form (i.e. evergreen cavo-convex).
winter hardy shrubs). flaccid Weak and lax, lacking firmness.
erose Irregularly toothed, as if gnawed; here often flexuous (Latin flexuosus) Bent alternately in op-
used in conjunction with denticulate. posite directions, in ‘zigzag’ fashion.
eutrophic Adjective referring to an environment flushing (of leaves) The more or less abrupt un-
(usually a body of water) rich in dissolved nu- folding from winter buds or just growth of new
trients available to plants. foliage leaves. In tropical trees, these young
evolution Gradual change (as contrasted with rev- leaves are often coloured pink or red, or a dif-
olution). In biology the term is mostly used in a ferent green from leaves that have been exposed
Darwinian sense and refers to (gradual) change longer.
of the genetic makeup of individuals in popu- foliate Leaf-like, i.e. shaped like a leaf as in a book
lations, primarily through genetic mutation, (Latin folio). Leaves of plants (conifers) can
separation (isolation) and natural selection, re- have shapes different from foliate, such as ac-
sulting in new species. The overall tendency is icular.
divergence, interrupted by extinction, causing foliola (Latin sing. foliolum) Little leaves or leaf-
major change over geological time. lets. A term used to describe two bract-like
exfoliating Here: the dead outer layers of bark appendages subtending the receptacle in seed
coming off in scales, sheets, or papery flakes. cones of the genus Podocarpus.
frondose Resembling a pinnately compound fern present. For Ginkgo there is abundant evidence
leaf (frond). of other species in the fossil record, for other
fusiform Spindle-shaped; tapering at both ends. genera with one extant species it is possible that
fynbos A vegetation formation occurring in the these will be found in future. [From J. R. Gregg
Cape Province of South Africa, characterized (1954). The Language of Taxonomy. Columbia
by a low, open structure and the abundance University Press, New York.]
of ericoid, proteoid, restioid and geophyte life gymnosperms Non-flowering seed plants with
forms on nutrient-poor soils. ovules directly exposed to air (‘naked’); the
gametophyte The generation (in spore-repro- seeds may be exposed also but are usually
ducing plants) that bears the sex organs; in enclosed within a cone or by an aril or epi­
gymnosperms strongly reduced to the micro­ matium; neither is technically a fruit (compare 1039
gametophyte in the pollen grain and the mega­ angiosperms).
gametophyte in an ovule or seed; the latter habit (Latin habitus = condition, character) Char-
refers to the tissue that embeds the (female) acteristic mode of growth or occurrence.
archegonia within an ovule; after fertilization of habitat The place or type of site where a plant (or
the archegonia the ovule develops into a seed. animal) naturally or normally grows and lives
The megagametophyte then has the same func- and to the natural conditions of which it is
tion as the endosperm in seeds of angiosperms, adapted.
but it has a different origin (homology). hardpan A compact soil surface inhibiting perco-
gene A bit of DNA (sequence of nucleotides) cod- lation of rain water to deeper levels where tree
ing for a protein, or part of a protein. roots could take it up. Such soils are common
genome The total of genetic or hereditary infor- in (semi) arid regions.
mation present in the DNA of an organism in heartwood The inner section of the wood of a
the form of the sequence of its nucleotides. tree, with dead wood cells through which no
genotype The genetic constitution of an individu- water is transported.
al or a group of organisms. helical (Latin helix) Arranged or formed in an up-
GIS A geographic information system (GIS) in- ward spiral (see also spiral).
tegrates hardware, software, and data for cap- heterosis A marked capacity for rapid growth of-
turing, managing, analyzing, and displaying all ten shown by cross-bred animals or plants (‘hy-
forms of geographically referenced informa- brid vigour’).
tion. heterotypic (synonym) A synonym based on a
glabrescent Becoming glabrous with age. different type. Such synonymy results from the
glabrous Devoid of hairs of any kind. merging of different taxa into one by taxono-
glandular Functioning as a gland, i.e. secreting a mists (compare homotypic).
biochemical substance. hexagonal Shaped like a hexagon, i.e. a polygon
glaucous Covered with a blue-green or blue-white with six angles and six sides.
bloom (see also pruinose). hilum A mark on a seed left by its connection to
globose Nearly spherical (compare globular). the tissue from which it developed.
globular Spherical, round (compare globose). hirsute Covered with coarse or shaggy hairs.
glutinous Sticky, with the quality of glue. holotype (Latin holotypus) The one specimen (or
graft The artificial growing connection of two illustration) used or designated by an author
parts of different plants, involving a root stock as the nomenclatural type of a species or lesser
and the grafted cutting or scion with foliage. taxon at the time of establishing it.
The resulting ‘plant’ minus the rootstock is also homologous Referring to shared characters in
referred to as a graft. two or more species due to common descent, or
Gregg’s Paradox This paradox seems to violate the to modifications evolved from such commonly
hierarchical structure of biological classifica- inherited characters.
tion: Ginkgo = genus → Ginkgo biloba = species homonym A name of an organism spelled exact-
→ Ginkgo = Ginkgo biloba → genus = species. ly the same as the name of another organism.
The way out of this paradox is to postulate Homonyms within the jurisdictions of the sev-
extinct species and only one surviving to the eral codes of biological nomenclature are not
 allowed and the rule of priority dictates that and regulations governing the naming of plant
the name shall be applied to the earliest named (and fungal) taxa, decided at International Bo-
plant or fungus (in case of the Botanical Code) tanical Congresses held at six year intervals and
and that the other plant or fungus must be re- organised by the International Association for
named. Plant Taxonomy (IAPT), which organisation
homoplasy (homoplasious) A term for a character publishes these rules in the Botanical Code (at
that is not a shared derived character (synapo- the time of publication of this Handbook the
morphy) in a phylogenetic sense. Homo­ ‘Vienna Code’ of 2006). Its rulings are binding
plasious characters (or character states) appear in plant taxonomy.
in more than one clade of an inferred phylo­ igneous Referring to rock that was formed from
1040 genetic tree (cladogram) and their similarity molten material (magma) in the interior of the
may result from convergent or parallel evolu- earth; it may have solidified below as well as on
tion, rather than from common descent with the surface.
modification or transformation. imbricate Closely overlapping, as thatches or tiles
homotypic (synonym) A synonym based on the on a roof (compare tesselate and valvate).
same type. Such synonymy results either from incertae sedis (Latin = place uncertain) A heading
the strict application of the Botanical Code under which taxonomists place names of taxa
(ICBN) or from a change in rank of a particular for which the original description is unclear in
taxon. Only the latter procedure involves taxo- the context of present knowledge and of which
nomic decision making (compare heterotypic). no good original material exists to determine
hort. (Latin hortulanorum) Of gardeners (horti­ their identity.
culturally); here added to a name used in a indehiscent Generally: a fruit that does not open.
(botanic) garden, but not formally published. If In conifer seed cones: the scales remain con-
followed by “ex” and a name and reference, its nate, so the cones will not open to release the
subsequent formal publication is given. seeds.
hybrid A product of propagation between two indigenous Naturally occurring in an area or a
individuals belonging to different taxonomic country, without having been transported there
units; hybridization can be naturally or arti- (intentionally or unintentionally) by people.
ficially induced. Some species of conifers are inflorescence A branching system supporting
believed to be of hybrid origin; their botanical flowers but usually not leaves.
names are marked with a × sign. in litt. (Latin: in litteris) In a letter or written note.
hypocotyl Section of a seedling between radix Citations given with this adjective are written
(primary root) and cotyledons, developing into but unpublished sources.
the stem. in sicco (Latin = in a dried state) Preserved as a
hypodermis Layer of one or more rows of thick herbarium specimen, of which the character
walled cells, situated directly beneath the epi- states may markedly differ from living or fresh
dermis; it is discontinuous at the stomata and material.
otherwise continuous, intermittent or absent. in situ (Latin = inside [its] place) In the context
hypogeal germination The hypocotyl of the seed- of plant species conservation, any effort to con-
ling does not elongate, the cotyledons remain serve individuals by protecting them or their
infolded below the soil surface and only the habitat within the natural range of the species
true stem of the seedling emerges (compare is considered in situ conservation (compare ex
epigeal). situ).
hypostomatic Stomata (almost) restricted to the integument Enveloping layer of the seed bud or
abaxial side of the leaves, with two primary ovule, which is open on one side (micropyle)
bands separated by a midrib in most species; and, after fertilization, develops into the seed
on plagiotropic lateral shoots the stomata are coat.
facing downward by a twisting of the leaf bases intercalary growth Growth inserted between
or petioles (compare epistomatic). other parts; in the bract-scale complexes of
ICBN International Code of Botanical Nomen- the seed cones of Cupressaceae this occurs in-
clature. A set of internationally accepted rules side the bracts (i.e. between the dermal layers)
 e­ nlarging them in various ways to form the that ecosystem and make it function. Removal
cone scales. of the keystone species destroys that particular
internal (as of resin canals or ducts) Inwardly; ecosystem.
here a position of the resin canals in the leaves knot In sawn wood the remainder of a branch in
bordering the endodermis. the stem of a tree, visible as a darker, rounded
introgression The entry or introduction of genes bit of wood with a different grain.
from one gene complex (e.g. a species) into an- K-strategy Also known as “K-selection”; in plant
other. ecology, this refers to trees with relatively slow
invers-dorsal A position of the leaves whereby growth rates but persistence (longevity) in later
the abaxial (‘dorsal’) side is inversed by twist- phases of vegetation succession. Its opposite,
ing of the leaf basis or petiole to face downward “r-selection” applies to trees that colonize early 1041
(‘­below’). phases (pioneer species), grow rapidly but are
inverted A position turned about 90° from an ear- later outcompeted by the slower, longer living
lier position; inverted seeds have turned their trees.
apex back towards their attachment point. lacerate Appearing as if irregularly torn or cleft.
involucre (Latin involucrum) A whorl of bracts laciniate Cut into narrow lobes or segments.
subtending a flower cluster (as in the heads lamina (Latin = blade, leaf) Here mostly used
of Compositae); in gymnosperms the whorl of for relatively broad, flat leaves of conifers that
scales (bud scales) subtending the strobilus in would by most people not be described as nee-
some species. dles or scale leaves.
isotype (Latin isotypus) A duplicate specimen of laminar Of the leaf; also referring to organs homo­
the holotype. logous to leaves.
IUCN International Union for the Conservation laminated Divided into lobes (as of leaves).
of Nature = The World Conservation Union. A lanate Woolly.
non-governmental organisation (NGO) with lanceolate Of a shape reminiscent of a lance point,
both government and non-government (e.g. i.e. much longer than wide and with the great-
academic) membership dedicated to the con- est width below the middle; commonly used to
servation of natural resources, in particular describe the shape of a leaf.
biological species. Its Species Survival Com- laterals (lateral leaves) The paired, opposite scale
mission (SSC) generates and updates the IUCN leaves on a plagiotropic foliage branchlet (esp.
Red List of Threatened Species™; its categories in the family Cupressaceae) that are positioned
and criteria are used in this Handbook. laterally, they flank and often partly enclose the
juvenile leaves Here the type of leaves that ap- differently shaped facials on either side.
pears on seedlings, saplings, or immature trees, laterite (Latin later = brick) A chemically weath-
or sometimes on reiterations; these leaves are ered, earth-like type of rock rich in oxidized
distinct from adult leaves, but intermediate metals.
forms may occur. lauraceous With alternate entire leaves character-
karst A deeply eroded formation of marine lime- istic of the family Lauraceae.
stone, of which surfaces are partly dissolved by layering The rooting of lower branches of a tree
rain water containing CO2 and where acidic where they reach the soil; from these branches
groundwater dissolves caves; drainage in karst new plants may grow, which are for a time still
landscapes is largely underground. In tropical part of the old plant but eventually become in-
climates karst landscapes develop tower hills or dependent when the parent plant dies.
mountains. lectotype (Latin lectotypus) A specimen or illus-
kerangas ‘Heath forest’, i.e. a more or less stunt- tration, designated from the ‘original material’
ed (but sometimes tall), open type of forest as the nomenclatural type if no holotype was
growing on nutrient-poor, sandy (often nearly indicated at the time of publication, or if it is
white) soil in tropical lowlands of Malesia. missing, or if it is found to belong to more than
keystone species A species in an ecosystem upon one taxon.
which many species depend that characterize
 leg. ign. (Latin legitavit = collected by, ignotus = Massart’s model A growth model in tree architec-
unknown). The abbreviation is used here when ture with a monopodial, erect stem with apical
the collector of a herbarium specimen was not dominance and branches placed in (pseudo)
stated. whorls at regular intervals; these branch again
lenticels Usually more or less raised, often some- and all or most branching is directed in a more
what corky spots on bark, of different shapes or less horizontal plane (plagiotropic), by sup-
but often horizontally elongate to linear, where pression of all but the laterally placed vegeta-
gaseous exchange takes place. tive buds.
lenticular Shaped like a lentil, i.e. with sides as in median A position ‘in the middle’ or central part
a double-convex lens. of an organ, e.g. a leaf.
1042 Life Zone A major altitudinal zone, characterized megagametophyte In gymnosperms (conifers)
by specific plant species and communities (and the tissue inside the integument of an ovule or
to a lesser degree by specific animal species) in seed coat of a seed, in which the fertilized gam-
a mountain region; they are specifically recog- ete develops into an embryo.
nized and named in Canada and the USA. membranous Thin, rather soft and more or less
lignified (Latin lignum = wood) Converted into translucent.
wood; become woody. meristem A formative plant tissue usually made
lignite An organic deposit from plant remains in up of small cells, capable of dividing continu-
an intermediate phase of being transformed ally and giving rise to similar or differentiating
from peat to bituminous black coal. cells.
lignotuber An outgrowth or swelling on the basal mesa (Spanish = table) A flat topped mountain
root system of a tree, forming excess wood and resulting from processes of erosion in ancient
capable of initiating new aerial stems with foli- sandstone or other sedimentary formations.
age and reproductive organs. mesic Characterized by or relating to a moderate
ligulate Tongue-shaped; also (in description of amount of moisture (see also mesophytic).
leaves) strap-shaped, resembling a ligulate mesophyll Leaf tissue, consisting of spongy pa-
flower in Compositae. renchyma and palisade parenchyma (the lat-
ligule (Latin ligula = small tongue) In the seed ter type not in all species) and filling the leaf
cone scale of Araucaria the apical free part of between the hypodermis and the endodermis.
the seed scale, which is for the most part fused mesophytic Growing under moist conditions; also
with the bract. of plants or plant communities growing under
linear An elongated, narrow, flattened shape with medium conditions of moisture, without (pro-
parallel or nearly parallel sides for all or most longated) periods of draught (see also mesic).
of its length. mesothermal Characterized by a moderate (cool)
lithosol An azonal shallow soil consisting of im- temperature without pronounced daily or sea-
perfectly weathered rock-fragments. sonal fluctuations.
Ma Million years ago. metamorphic Of a rock type that has been sub-
maquis A thick, scrubby (underbrush) vegetation stantially altered due to high pressure and/or
of predominantly xerophytes (e.g. sclerophyl- heat generated by earth crust movements.
lous shrubs) characteristic of Mediterranean micropyle The opening in an ovule through which
shores and mountain slopes. [In New Caledo- pollen enters, in most conifers with the aid of a
nia, similar vegetation (but with very different pollination drop of fluid.
plant species) growing on ultramafic soil rich microsporangium (plural: microsporangia) A
in metals is called ‘maquis minier’, as these sub- sac-­like structure that contains the micro-
strates are often mined to extract nickel.] spores, it is dehiscent so it can release the ripe
Malesia A floristic region of mainly islands in spores. In conifers this is usually termed pollen
SE Asia, defined (mapped) in Flora Malesiana sac and it contains pollen.
as extending from the Isthmus of Kra on the microsporophyll A (modified) leaf-like struc-
Malay Peninsula to the end of the main archi- ture bearing the pollen sacs containing pollen
pelago of the Solomon Islands. The serial Flora or male gametes (in gymnosperms); they are
Malesiana describes the plants of this region. borne on the male strobilus or pollen cone.
molecular analysis In the present context a phy- mucronate Having a mucron, i.e. a narrow, abrupt
logenetic analysis using DNA sequence data as point which is usually only a continuation of
characters and cladistic methodology. the midrib (or midvein).
monoecious In gymnosperms this pertains to fe- muskeg (word of Algonquin origin) A Sphag­
male and male reproductive organs, commonly num-built bog of northern latitudes in North
cones, being present on the same plant. Plants America, often with scattered and stunted tree
with only one sex present are dioecious; both growth.
conditions occur in conifers, with some genera mutant An organism that obtained a genetic char-
(or families) having one condition in all spe- acter change (mutation) not present in other
cies, but others more variable. individuals of its species.
monograph In biology the comprehensive taxo- mutualism An ecological (or behavioral) relation- 1043
nomic description of a natural group of species, ship between two species that is beneficial to
usually a genus or a family, including all aspects both.
pertinent to the systematics and classification mycelian Of the mycelium, the threadlike net-
of that group. work of hyphae of fungi.
monomorphic Of single form or shape (= 1 char- mycelium A mass of filamentous hyphae, usually
acter state); all similar organs (e.g. leaves) have underground or in wood, which forms the veg-
the same shape (compare dimorphic and poly- etative portion of a fungus.
morphic). mycorrhiza The thread-like hyphae of certain
monophyletic Adjective referring to a group of fungi that form intricate connections with the
taxa; originally (Ernst Haeckel 1866) used in roots of plants and live in symbiosis with these
the sense of sharing a unique common ances- plants.
tor, but now almost universally used in the cla- mycropyle The small opening at the apex of an
distic sense (Willi Hennig 1950, 1966), in which ovule, usually shaped somewhat like the neck
it refers to an ancestor and all its descendants. of a flask, through which pollen can enter.
[A more accurate term would be holophyletic.] natural group A group of taxa that is based on a
Groups that do not include all descendants are hypothesis of common descent from a nearest
then described as paraphyletic. ancestor.
monopodial Having a single stem or trunk (bole), naviculate (Latin navicula = a boat) Boat shaped,
usually growing more or less straight up from referring to the shape of open boats with nar-
the ground. rowing bow and stern and a more or less keeled
monospecific Referring to a taxon of higher rank bottom.
containing a single species (see also mono­ neotype (Latin neotypus) A specimen chosen and
typic). designated as the type of a taxon in a situation
monotypic Referring to a taxon containing only a where no original material is extant.
single taxon of lower rank, such as a genus with niveous white Snow-white, without any tinge of
only one species (see also monospecific). different (underlying) colours.
montane Growing in, or being, the biogeographic nom. cons. (Latin nomen conservandum = re-
zone that is made up of relatively moist cool tained name) A botanical name invalid under
upland slopes below timberline in mountain- the rules of the Botanical Code (ICBN), but re-
ous areas. tained to avoid disadvantageous changes in the
morphology The detailed study of form; in sys- nomenclature by incorporation in the append-
tematic research of taxa this involves investi- ed lists of nomina conservanda (et rejicienda)
gations into anatomy and development as well published in the ICBN.
as comparison among individuals and taxa in nom. illeg. (Latin nomen illegitimum = illegitimate
order to understand characters and their states. name) A botanical name invalid under the
mossy forest A high montane to subalpine forest rules of the Botanical Code (ICBN), as speci-
type in the tropics of SE Asia, characterized by a fied under Art. 6.4 (Vienna Code, 2006).
low canopy of flat-topped crowns of trees, some nom. inval. (Latin nomen invalidum = invalid
of them emergent, and draped with great quan- name) A botanical name that is not validly
tities of epiphytic mosses, lichens and ferns. published under the rules of the Botanical
 Code (ICBN). Unlike an illegitimate name, it in obovate Inversely egg-shaped in outline (2-di-
fact does not exist for nomenclatural purposes mensional).
unless and until it has been validated under the obovoid Inversely egg-shaped (3-dimensional,
rules. compare pyriform).
nom. nud. (Latin nomen nudum) A botanical obtrullate Inversely trullate, i.e. shaped like a
name without a formal description and there- bricklayer’s trowel with the two short sides at
fore invalidly published under the rules of the the apex.
Botanical Code (ICBN). obtuse Blunt, with a more or less rounded apex.
nom. ut. rej. (Latin nomen utique rejiciendum = oligotraphent Adjective of a taxon, which pre-
rejected name in any case) A botanical name dominantly grows in an oligotrophic environ-
1044 ruled as rejected under Art. 56 of the Botanical ment.
Code (ICBN), listed in Appendix V and not to oligotrophic Adjective of an environment (usually
be used in any case. a body of water), which is poor in dissolved nu-
nothospecies (nothotaxon) A species (or taxon) trients available to plants (compare eutrophic).
which originated from the hybridization of two ontogenetically (ontogeny) Referring to the proc-
other species (or taxa) and is found to form ess of development or growth of an organism or
populations beyond the F1 generation with dis- its parts from fertilized egg to adult.
tinct and more or less stable characters. op. cit. (Latin opus citatus) Work cited. Referring
nucleotide Any of the unit building blocks or bas- here to the publication cited with a name of a
es of DNA and RNA, commonly represented by taxon under the species to which the text ap-
the letters A (adenine), C (cytosine), G (gua- plies; instead of to the list of references at the
nine) and T (thymine); in RNA U (uracil) re- end of the Handbook.
places T. In the DNA chain, A links with T and orbicular Approximately circular in outline.
C with G, forming sequences of nucleotides. orthotropic Growing of the stem (sometimes the
numerical taxonomy (phenetics) Taxonomy de- branches) in a vertical or near vertical direction,
rived from the statistical analysis of variable as opposed to the plagiotropic lateral branches
data, i.e. the calculation of phenetic similarity growing in a (near) horizontal direction.
between two taxa based on as many measur- ovate Egg-shaped in outline (mostly for plane
able variables (character variations) as possible, shapes).
usually by applying one of many available ‘clus- ovoid Egg-shaped.
tering’ algorithms and statistical probability ovulate Referring to or having ovules.
tests. Greater overall similarity is assumed to ovule Seed bud, in gymnosperms (conifers) in-
mean closer relationship, thus directly inform- cluding the embryonic stages of a seed up to the
ing the classification. event of fertilization, in which it is exposed to
nutrient In the context of plants any of the basic air and develops specialized features to capture
elements such as nitrogen (N), potassium (K) pollen.
and phosphor (P) necessary for growth and ovuliferous Bearing the ovules (Latin ovulum);
physiological processes. which after pollination and fertilization be-
n.v. (Latin non vidi) I have not seen it. Added to come the seeds.
the citation of a (type) specimen one knows palaeobotany A branch of botany (or geology)
exists, but has not been able to see. Viewing a concerned with the study of fossil plants or
specimen can include seeing an image of it, e.g. their remains.
on the Internet. palaeontology The science or study of fossil or-
obcordate Inversely heart-shaped. ganisms.
obdeltoid Shaped like an inversed capital Greek palisade cells Perpendicular elongated parenchy-
letter ‘delta’ (triangle). ma cells in the mesophyll near the surface of
oblanceolate Considerably longer than broad, in- leaves.
versely lance-shaped (the widest part above the palynology The science or study of pollen and
middle, compare lanceolate). spores, both of recent and fossil plants.
oblong Considerably longer than broad, with paniculate Arranged in a panicle, i.e. a compound,
nearly parallel sides. branching raceme.
papilliform Shaped like a nipple (Latin papilla). perular scales The basal scales of buds which re-
parallelism In an evolutionary context, the devel- main at the base of shoots or cones some time
opment of a similar character in two or more after elongation or maturity.
lineages which do not share a nearest common petiolate Having a leaf-stalk (petiole); also: shaped
ancestor; the character has therefore evolved like a petiole.
more than once and independently. phenotypic (phenotype) Relating to the anatomi-
paraphyletic Adjective referring to a group of taxa cal and morphological appearance (characters)
that share a nearest common ancestor but, as of an organism, in particular those determined
a group, does not include all of that ancestor’s by its genes (DNA, genotype) and a concomi-
descendants (see also under monophyletic). tant ontogeny.
paratype (Latin paratypus) A specimen cited in photosynthesis Synthesis of chemical compounds 1045
the protologue that is neither the holotype nor with the aid of radiant energy (light), resulting
an isotype, nor one of the syntypes if two or in the formation of carbohydrates (assimilates)
more specimens were simultaneously desig- from carbon dioxide and water in chlorophyll-
nated as types. containing plant leaves (see also under chloro-
parenchyma A tissue of higher plants composed phyll).
of thin-walled cells; especially such tissue as the phyletic Referring to both common ancestry re-
pith and mesophyll. lationships (phylogeny) and ancestor/descend-
passim (Latin passus = scattered) After a page ant relationships (diversification of characters
number, indicating that the name (subject) /character states through a hypothetical evo-
subsequently appeared on several other pages. lutionary lineage resolved in the phylogeny)
pathogenic Referring to a pathogen, an agent or (compare phylogenetic).
organism that causes disease. phylloclade (phyllode) (Greek phyllos = leaf,
pectinate Arranged as the leaflets of a compound, clados = branch) A leaf-like branch; in the ge-
pinnate leaf; with the needles (leaves) arranged nus Phyllocladus true leaves are rudimentary
on both sides of the shoot, spreading sideways and foliage branchlets are strongly flattened,
and more or less in a plane (compare pinnate). functioning as leaves.
pedicellate With a small flower stalk (Latin pedi­ phyllotaxis The arrangement of leaves on a shoot
cellus); here used to describe the abruptly nar- or stem and their position in relation to each
rowed base of seed scales. other.
peduncle The stalk of a flower or inflorescence phylogenetic Referring to common ancestry re-
(Latin pedunculus); usage of this term to desig- lationships between extant species or taxa of
nate the stalk of a conifer (female) cone is mor- higher ranks (see phylogeny and compare
phologically incorrect insofar as such a cone is phyletic).
not an inflorescence, but a shoot with ovuli­ phylogeny A hypothesis of common ancestry rela-
ferous scales, but has been common practice tionships of taxa, usually depicted as a branch-
and is maintained here for convenience (and ing diagram or ‘tree’ (‘Tree of Life’) showing
likewise used for the stalk of male strobili). branching relationships from a hypothetical
peltate Shaped more or less like a shield, with an common ancestor to multiple descendants.
attachment more or less close to the centre. Cladograms, although really only showing re-
pendant Hanging down (nearly) vertically (see lationships of characters and character states,
also pendulous). are usually interpreted as phylogenies by in-
pendulous Hanging down (nearly) vertically (see serting taxa on the (terminal) branches, infer-
also pendant). ring their phylogenetic relationships from the
permafrost A permanently frozen layer of sedi- branching pattern (topology) of the cladogram.
ments or soil, of variable thickness, formed in phytogeography The study of natural plant (usu-
cold regions; of this layer the thin upper part on ally species) distribution and its causes.
the surface thaws in summer. pilose Having straight, soft and spreading hairs.
pentagonal Shaped like a pentagon, i.e. a polygon pinetum (plural: pineta) A park or garden dedi-
with five angles and five sides. cated to a living collection of planted conifers.
 pinnate Having the arrangement of a feather, with primary branches The branches that appear more
a single rachis from which leaflets arise on ­either or less rhythmically during the growth of a tree,
side (compare pectinate). from first to highest order. Secondary branches
plagiotropic Growing of (lateral) branches in a are the result of reiteration. The distinction be-
horizontal or near horizontal direction, away tween the two categories becomes increasingly
from the (near) vertical (orthotropic) stem or blurred and often difficult to determine in old
first order branches. trees, in which much of the main branches may
planation The transformation through evolution represent reiteration.
of a more or less terete shape of something in primary leaves The first leaves of a young plant,
an ancestor into one that becomes flattened and other than the cotyledons. In some genera (or
1046 wider in its descendants. species) these leaves differ markedly from later,
plate tectonics The formation and dynamics of adult types of leaves, e.g. in Pinus (cataphylls),
the plates into which the crust of the Earth is in others the differences are gradual, e.g. in
divided. Podo­carpus (compare secondary leaves).
podzolic (podzolized, Russian ‘podzol’ = ‘under primary (bud) scales The outer scales, which are
ashes’) Referring to an infertile, sandy and often much longer than the inner (secondary)
acidic soil with minerals leached from its sur- scales in foliage branches of the genus Podo­
face layers and deposited in a lower stratum. carpus.
The leached part of the profile is ash-grey. prismatic Shaped like a prism, i.e. with polygonal
pollen (singular and plural) The (near) micro- faces lying in parallel planes.
scopic unit or ‘grain’ in seed plants that con- proliferation (in conifers) The development of a
tains the male gamete enclosed by a hard and foliage shoot from a normally determinate or-
usually multi-layered wall and is dispersed to gan, e.g. a seed cone.
meet an ovule for fertilization. protologue (Greek protos =first; logos = discourse)
pollen sac (microsporangium) A variously shaped Everything associated with a name at its first
sac-like vessel containing pollen and which is valid publication, i.e. diagnosis, description,
borne on the microsporophyll of a pollen cone illustrations, references, synonymy, geographi-
in conifers. cal data, citation of specimens, discussion, and
pollination The transfer of pollen from a stamen comments.
(in angiosperms) to an ovule; in gymnosperms provenance Information about the source of a
the pollen descends directly on the nucellus plant or its seed used in forestry or horticulture.
and germinates; the pollen tube begins to grow proximal Occurring nearest the base or axis
and penetrates the nucellus. (compare distal).
polymorphic Of various forms or shapes; similar pruinose Covered with a coarse, whitish, waxy
organs on the same plant appear in > 2 char- bloom, more prominent than if glaucous.
acter states, e.g. variable leaf forms (compare pseudo-whorl An arrangement that is seemingly
dimorphic and monomorphic). forming a whorl, but is in fact helical on a very
polyphyletic Referring to a group of taxa that do short axis.
not share a nearest common ancestor. puberulent Minutely pubescent, i.e. covered with
polytomy In a cladogram referring to lineages very small, soft hairs.
(clades) with unresolved relationships; such pubescent Covered with soft, short hairs.
lineages all arise from a single node. pulverulent Being or looking dusty or powdery
population The sum total of the individuals of a (from almost invisible short hairs).
species (or lower taxon) which inhabit a certain pulvinus (plural: pulvini) Small, peg-like projec-
area and are expected to interbreed (exchange tion on the shoot supporting the leaf (needle)
genes). in several genera especially of Pinaceae.
p.p. (Latin pro parte = in part) adjective to a name pungent Ending in a rigid, sharp point or prickle.
indicating that only a certain part (including pustulate Having pustules, i.e. low projections like
or excluding the type) of the original material a blister or pimple.
mentioned in the protologue is (to be) con- pyriform Shaped like a pear, i.e. with the broadest
nected with that name. part near the apex (compare obovoid).
quadrate In fours, here pertaining to the phyllo- revolute Rolled downward or backward.
taxis, with four leaves originating at exactly the rheophyte A plant that completes its life cycle in
same distance on the shoot. If one of the op- streams, but is not aquatic; a rheophyte germi-
posite pairs is slightly higer, the arrangement is nates out of water when the stream falls tem-
decussate. porarily dry or recedes from normally higher
quadrate-rhombic Shaped between quadrangular levels.
and rhombic (compare rhombic). rhombic Having the shape of a rhombus, i.e. an
raceme An inflorescence with the oldest flowers equilateral parallelogram usually having ob-
(pollen cones in conifers) the most proximal lique angles (compare rhomboid).
and a potentially continuously growing apex. rhombic-orbicular Shaped between rhombic and
racemose In the form of a raceme. orbicular. 1047
rachis The axis (homologous to a shoot) of the rhomboid shaped like a parallelogram in which
cone of a conifer, from which bracts, micro- the angles are oblique and adjacent sides are
sporophylls or seed scales arise. unequal (compare rhombic).
Rauh’s model A growth model in tree architec- riparian Growing on the banks of rivers, subject
ture with a monopodial, erect stem with apical to flooding, but not permanently in water.
dominance and branches placed in (pseudo-) RNA Ribonucleic acid. In its various forms, RNA
whorls at regular intervals; these and secondary acts as the intermediary by which the heredi-
branches are assurgent and by erecting them- tary code of DNA is converted into proteins
selves (becoming orthotropic) cause repetition (compare DNA).
of the first order branching in (pseudo-)whorls. rostrate Shaped like a beak (Latin rostrum).
receptacle A fleshy or succulent structure sub- ruderal Referring to plants or vegetation growing
tending a free standing seed in Podocarpus and in and adapted to continuously or repeatedly
some other genera of the Podocarpaceae; it is disturbed sites (dynamic habitats).
formed from all remnants of the seed cone after saccate With bladders (Latin sacci); some types of
fertilisation of usually a single egg (ovule) de- pollen in conifers have 2 (rarely more) hollow
veloping into the seed. bladders, presumably to aid buoyancy.
recurved Curved backward (compare decurved). sapwood The outer section of the wood of a tree,
refugium (plural: refugia) A geographical area with ‘living’ wood cells through which water is
into which one or more species have retreated transported.
(or where they remained) from a much wider scarious Thin, dry and membranous, not green
distribution in the past. (herbaceous).
reiteration (Latin reiterare = to say or do repeat- scion A vegetative shoot cut from a plant and
edly) The secondary initiation of branching caused to produce roots or grafted onto a dif-
from a primary branching system (or from the ferent rootstock.
trunk) of a tree, activating dormant buds, usu- sclereids (Greek skleros = hard) Inclusions in the
ally as a response to damage. mesophyll or below the surface of leaves con-
rendzina A dark greyish brown intrazonal soil sisting of ‘stone cells’, i.e. dense and hard bodies
developed in usually grassy regions of high to often with a random position and distribution.
moderate humidity from soft calcareous marl sclerified Hardened by transformation or addi-
or chalk. tion of cells with lignified walls.
reniform Kidney-shaped; much wider than long. sclerophyllous With hard leaves; leaves with a
repand With an undulating or wavy margin. tough texture, mainly from dermal cells with
resin duct (resin canal) A tubular intercellular lignified walls.
space, especially in gymnosperms, that is lined secondary leaves Leaves of the ‘adult’ type; in con-
with cells which secrete resin. ifers these may appear soon on young plants, or
resin vesicle A small cavity or bladder filled with they may be delayed for much longer and they
resin. are the forms that are usually associated with
reticulate Forming a network of connections or fertile branches and/or appear in the canopy
relationships between distinct entities. of mature trees or shrubs (compare primary
retuse Notched shallowly at a rounded apex. leaves).
 seed scale Appendage in a conifer seed cone (situ- sister (group or taxon). In cladistic terminology
ated in the axil of a bract) bearing one or more a group or a taxon that is a clade next to an-
seeds. other clade, of which it is the sister clade; with-
sensu lato (Latin = in a broad sense) In taxonomy in a particular cladogram these two are more
it follows a taxon name to indicate that it in- closely related to each other phylogenetically
cludes another taxon at the same rank, which than they are to any other clades (groups) rep-
some may recognize as distinct (abbreviated as resented.
s.l.). The opposite is sensu stricto. s. l. (Latin sensu lato) In a broad sense; used in ad-
sensu stricto (Latin – in a strict or narrow sense). dition to a taxonomic name to differentiate it
In taxonomy it follows a taxon name to indicate from the use of that name in a strict sense (see
1048 that it excludes another taxon at the same rank, sensu lato).
which some may consider as synonymous (ab- s.n. (Latin sine numero) Without a number. If col-
breviated as s. str.). The opposite is sensu lato. lectors of botanical specimens did not give a
septal (Latin septum = enclosure) Here a position number to them, such collections are cited with
of the resin canals in the leaves wedged between the name of the collector and ‘s.n.’ immediately
the endodermis (enclosing the inner vascular following.
cylinder) and the hypodermis. s. str. (Latin sensu stricto) In a strict or narrow
sequencing (of DNA) A technique using bio- sense (see sensu stricto).
chemical methods to determine the sequence spathulate Spoon-shaped.
of the four nucleotides (the genetic code) in a speciation To differentiate into new biological
particular section of the DNA strand of an or- species through evolution, involving genetic
ganism. These sequences are copied through separation of populations and divergence of
generations, but small changes (mutations) oc- characters (character states) through time.
cur from time to time, which are again copied species complex Term used to describe a group of
and so serve as markers of heredity. They can be closely related species, within which group spe-
identified and scored as derived characters for a ciation takes place (variation leading to genetic
cladistic analysis. separation) and/or which is often characterized
sere A successional sequence of plant associations by hybrids (occurrence of hybrid-swarms, re-
or communities that replace each other over ticulate relationships between taxa).
time at a given site, usually going from pioneer speciose Counting or having many species.
stages to later stages in which an ecological spicate As in a spike, spike-like.
equilibrium is reached. The stages are called spike An inflorescence consisting of an axis and
seral to emphasize their transitory nature. numerous flowers (pollen cones in conifers) ar-
serotinous (Latin sero = late) Late to appear or ranged on it (see also raceme).
flower; in conifer seed cones pertaining to a de- spinescent Shaped like a spine or thorn (compare
layed opening, usually associated with persist- pungent).
ence on branches; both are adaptations to fire. spiral Arranged or shaped in an outward going,
serrate Saw-toothed, the sharp teeth pointing for- circular fashion. On shoots, this will be an up-
ward. ward spiral and is more accurately termed heli-
serrulate Finely serrate. cal, but “spirally arranged leaves” is often used,
sessile (Latin sessilis = sitting, attached) Without including in this book.
a peduncle or stalk, or in the case of conifer sporophyll Literally a leaf bearing spores (as in
cones, with a very short one which is invisible ferns); in gymnosperms it refers to leaf-like ap-
under the basal scales so that the cone appears pendages bearing male (= microsporophyll) or
to be stalkless. female (= macrosporophyll) reproductive or-
sister group A clade in a cladogram that is near- gans: pollen (in pollen sacs) or ovules.
est in relationship to another clade, or a taxon squarrose With stiff or rigid branches or protru-
represented by such a clade. The term usually sions.
(but not necessarily) refers to the clade below stele An axial cylinder of tissue in which the vas-
the next one in a cladogram. cular tissue is developed; its outer layer of cells
forms the endodermis.
stomata (sing. stoma) Breathing pores or aper- by its descendants; it could be lost subsequently
tures in the epidermis, surrounded by two in some descendants and transformed into a
guard cells (and subsidiary cells), leading into different character in others (see also homo­
an intercellular space communicating with the logous).
internal tissue. taiga (Russian = forest) Northern coniferous for-
striated Marked with (longitudinal) lines or est between tundra in the north and steppe in
streaks. the south, particularly in Asia.
strobilus (Greek strobilos = pine cone; plural: talus A slope formed especially by accumulation
strobili) The technical term for the reproduc- of rock debris from rock formations above it.
tive unit of a gymnosperm that bears male or taxon (plural: taxa) Any group of organisms that
female organs, regardless of whether it forms a is recognized at any of the ranks of classifica- 1049
distinct cone or not. tion used by taxonomists, such as family, genus,
subalpine Referring to, or growing in the biogeo- species.
graphic zone between the montane zone and taxonomy The science of classification of organ-
the tree limit in high mountains; it is the alti- isms and the identification and naming of taxa.
tude of the highest growing tree species in a TDWG International Working Group on Taxo-
given area. nomic Databases for Plant Sciences (TDWG).
substrate The base on which an organism lives; for The codes used in this Handbook are for po-
most vascular plants this is the soil. litically defined geographical areas (countries,
subtending Occupying an adjacent and usually provinces etc.) according to the World Geo-
lower position to and often (partly) enclosing graphical Scheme for Recording Plant Distri-
an organ. butions, compiled by R. K. Brummitt (2001).
subterete Almost terete, the shoot may be very See with this reference for access.
slightly angular or ridged. terete Approximately cylindrical.
subulate Awl-shaped; linear and tapering to a fine terminal Referring to the position of an organ or
point. structure at the tip of a shoot.
succession In ecology, the gradual and successive ternate Occurring in threes; leaves are ternate
replacement of plant species by others in one only if all 3 leaves originate at exactly the same
locality due to development of the vegetation distance on the shoot, otherwise they are more
from a pioneer phase to a climax phase. likely to be alternate.
succulent Of a plant: having fleshy tissues adapted terpenoid (German: Turpentine) Referring to
to conserve and store moisture (like the cac- chemical compounds based on a five-carbon
tuses or cacti). atom structure and present in conifer resins,
symbiosis A mutually beneficial physiological re- also known as terpenes.
lationship between two or more different spe- tesselate elements or pieces laid closely together as
cies, with give and take in equilibrium in the in a mosaic or like paving stones, joined only at
sense of an evolutionary stable strategy (ESS). the margins (compare imbricate).
sympatric Occurring in (generally) the same area tetragonal Having four sides or faces derived from
or with partly overlapping areas, but without a quadrangular base.
loss of genetic identity by interbreeding. tetrastichous Arranged to forming 4 apparent
syntype (Latin syntypus) Any specimen cited in rows from a helical leaf attachment on the
the protologue when no holotype was desig- shoot.
nated, or any of two or more specimens simul- tomentose Densely covered with short, woolly
taneously designated as types. hairs.
systematics The science of the diversity of and the tracheid A dead cell in the wood of gymnosperms
relationships between taxa based on evolution- characterized by lateral pits in the cell walls
ary principles, including (or sometimes seen as connecting the lumen of one tracheid with that
synonymous with) taxonomy. of another, allowing fluid transport; tracheids
synapomorphy (plural: synapomorphies) Shared are structurally distinct from the equivalent
derived character(s); i.e. a character originated cells in angiosperms (vessels).
in a common ancestor and inherited unaltered
 translucent Of a substance or thing with a colour, ultramafic Referring to rock or soil poor in sili-
density, or thickness that allows some light to ca, but extremely rich in iron and magnesium
penetrate, but insufficient to be seen through minerals and with a high pH value (also ultra-
(compare transparent). basic).
transparent Of a substance or thing with a colour, umbellate Borne in umbels, i.e. a (flower) cluster
density, or thickness that allows it to be seen in which the pedicels or stalks (branches) arise
through (compare translucent). from a common point.
transverse Made at right angles to the anterior- umbilicate Depressed like a navel.
posterior axis of a body or structure; also set umbo (orig. Latin = the boss of a shield) In pine
crosswise. cones it is a prominence on the apophysis of
1050 transversely rhombic Having the shape of a the scales, often armed with a prickle or spine.
rhombus (see rhombic), which is transversely undulate Having a wavy surface, edge, or mark-
widened (see transverse). ings (see also repand).
triangulate Appearing as a triangle but not really utriculate Inflated, bladder-like.
shaped so. valvate Parts connecting or touching with their
tridentate Three-toothed; in the bract scales of edges (as in the two shells of an oyster), not
cones of Pinaceae with a short, pointed central overlapping (compare imbricate).
lobe flanked by two short, pointed lateral lobes variegated Variably coloured in foliage or leaves,
(see also trilobate). usually occurring in cultivars from a partial
trigonal Having three sides or faces derived from deficiency in the amount of chlorophyll in the
a triangular base. leaves, causing a yellowish hue contrasting with
trilobate Three-lobed; in the bract scales of cones normal green.
of Pinaceae with a long or rounded central lobe vascular bundle A unit of the vascular system of
flanked by two long or rounded lateral lobes a vascular plant, consisting of vessels and sieve
(see also tridentate). tubes, together with parenchyma cells and
trimerous In threes; i.e. made up of three separate, ­fibres; it has a function in transportation of
free parts (compare tripartite). ­water, nutrients and assimilates.
tripartite Made up of three separate but connect- vegetative bud (vegetative apex) A bud which
ed parts (compare trimerous). gives rise to the vegetative organs, i.e. the shoot,
trisaccate (in pollen) Having three sacci or air dwarf shoots and needles or leaves. In the de-
bladders (compare bisaccate). scriptions of this Handbook usually the ter-
trullate Shaped like a bricklayer’s trowel. minal buds at the end of branches are the only
truncate Ending abruptly, as if cut off transversely ones described (see also winter bud).
(but usually with rounded corners). venation The number and distribution (pattern)
tussock In grassland, clumps formed by the con- of veins in a leaf.
stituent grasses that are raised above the gen- verrucose Covered in numerous small, wart-like
eral surface, such grassland is very uneven as elevations (verrucae).
a result. vessel (in wood) A cell in the wood of angiosperms
type (type specimen) A specimen designated as (and some gymnosperms) with closed lateral
the type of the name of a species or taxon with walls and distal openings allowing fluid trans-
lower rank, fixing the application of that name port (compare tracheid).
by including its characters. The circumscrip- vicariance (Latin vicarius = substitute) In bio­
tion of a taxon can be widened or narrowed, geography referring to the distribution of taxa
but always must include that of the type speci- explained by the history of the separation of
men if it is to retain its name. Similar principles the areas in which these taxa occur. The term
apply at higher ranks, where e.g. a species name originally referred to a method of analysis in
becomes the ‘type’ of a genus, but in fact refers which area cladograms were substituted for
to the type specimen of that species name. taxon cladograms. This concept is intimately
ultrabasic Referring to rock or soil with a high pH ­connected with cladistic methodology, un-
rich in metallic minerals (also ultramafic). der the assumption that distribution patterns
 and speciation are caused by the same historic the young shoot of which the internodes are
events. It ignores (later) distribution of taxa not yet elongated; it will start growing at the
caused by dispersal of organisms. beginning of the spring season (see also veg-
vicariant (Latin vicarius = substitute) A taxon etative bud).
occupying a similar niche as a different, but xeromorphic Structurally adapted for life and
related taxon occurring in a separate area growth with a limited water supply, especially a
(vicariants are allopatric). The two taxa are morphology limiting transpiration (in gymno-
inferred to have derived from an (extinct) com- sperms) or providing for the storage of water;
mon ancestor and their separation the result of such plants are xerophytes.
the emergence of an ecological barrier between xerophyte A plant structurally adapted for life and
them. growth with a limited water supply; usually by 1051
whip shoot A relatively fast growing, leading fo- means of xeromorphic adaptations that allow
liage branchlet; due to its faster growth scale water storage and ensure limitation of transpi-
leaves, which grow with it, are longer and often ration.
also wider than those on short, lateral branch- zoochory The dispersal of plant propagules (e.g.
lets. seeds) by animals.
winter bud A terminal bud, which in conifers is
covered with scales or cataphylls which can be
resinous or non-resinous, and which contains
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LISTS OF ILLUSTRATIONS

Map showing the global distribution of conifers 21

List of line drawings (plates)

1065
1. Abies bracteata 67 22. Nothotsuga longibracteata 534
2. Abies mariesii 101 23. Picea alcoquiana 559
3. Abies nephrolepis 104 24. Picea neoveitchii 588
4. Abies nordmanniana 106 25. Picea torano 603
5. Agathis borneensis 149 26. Pinus arizonica var. arizonica 622
6. Araucaria angustifolia 189 27. Pinus canariensis 645
7. Araucaria scopulorum 213 28. Pinus dalatensis 660
8. Callitris monticola 233 29. Pinus flexilis 675
9. Callitris verrucosa 244 30. Pinus krempfii 694
10. Cedrus deodara 257 31. Pinus lambertiana 697
11. Chamaecyparis obtusa var. obtusa 279 32. Pinus pinceana 736
12. Cupressus macnabiana 310 33. Pinus pseudostrobus 743
13. Dacrydium beccarii 342 34. Pinus torreyana subsp. torreyana 775
14. Fokienia hodginsii 373 35. Podocarpus brassii 816
15. Juniperus drupacea 408 36. Podocarpus elatus 833
16. Juniperus excelsa subsp. polycarpos 413 37. Podocarpus lambertii 848
17. Juniperus monosperma 426 38. Podocarpus pendulifolius; P. salicifolius 878
18. Juniperus recurva var. recurva 443 39. Podocarpus steyermarkii 902
19. Keteleeria fortunei 477 40. Podocarpus urbanii 911
20. Larix potaninii 494 41. Tetraclinis articulata 986
21. Lepidothamnus laxifolius 501 42. Xanthocyparis vietnamensis 1031

List of colour photographs (figures)

The colour photographs have been grouped into six sections.

First section (Fig. 1–59) 177–184

1. Abies alba in the Forêt de la Joux, Jura Mts., 10. Abies koreana young seed cones
France 11. Abies magnifica in the Sierra Nevada, Califor-
2. Abies amabilis in Wenatchee N. F., Washington, nia, USA
USA 12. Abies nebrodensis seed cone
3. Abies bracteata in the Santa Lucia Mts., Califor- 13. Abies numidica tree with seed cones
nia, USA 14. Abies pinsapo helically arranged leaves
4. Abies cephalonica flushing leaves 15. Abies pinsapo seed cone
5. Abies delavayi var. delavayi seed cones 16. Abies procera seed cones (photo W. Milliken)
6. Abies delavayi var. delavayi seed cone 17. Abies recurvata var. ernestii seed cones (photo
7. Abies delavayi var. nukiangensis seed cones X. C. Zhang)
8. Abies homolepis pollen cones 18. Abies spectabilis mature seed cone
9. Abies kawakamii stand in Taroko N. P., Taiwan
 19. Abies squamata bark of young tree (photo P. de 38. Amentotaxus formosana tree in southern Tai-
Spoelberch) wan (photo C. N. Page)
20. Abies squamata bark of old tree (photo P. de 39. Amentotaxus formosana leaves underside
Spoelberch) 40. Amentotaxus yunnanensis in Ha Giang, Viet-
21. Acmopyle pancheri foliage and see cones nam (photo P. Cribb)
(© Bedge­bury Pinetum) 41. Araucaria araucana in Chile (photo M. Gard-
22. Actinostrobus arenarius along Hwy. 123, West- ner)
ern Australia 42. Araucaria araucana pollen cones
23. Actinostrobus arenarius seed cones 43. Araucaria araucana seed cones
24. Afrocarpus falcatus foliage and seeds (photo 44. Araucaria bernieri in New Caledonia
1066 D. Luscombe) 45. Araucaria bidwillii emergent trees in the Bunya
25. Afrocarpus gracilior tree in Ethiopia (photo Mts. Queensland, Australia
J. Grimshaw) 46. Araucaria bidwillii tree in the Bunya Mts.
26. Afrocarpus gracilior foliage and seeds (photo Queensland, Australia
Forest & Kim Starr) 47. Araucaria columnaris on the Île des Pins, New
27. Agathis australis in Trounson Park, North Is- Caledonia
land, New Zealand 48. Araucaria columnaris seedlings
28. Agathis australis The Tane Mahuta (Lord of the 49. Araucaria cunninghamii var. cunninghamii tree
Forest), New Zealand in the Bunya Mts. Queensland, Australia
29. Agathis australis seed cones 50. Araucaria heterophylla seed cones
30. Agathis borneensis leaves 51. Araucaria laubenfelsii foliage with pollen cones
31. Agathis kinabaluensis on the Mesilau River, Mt. 52. Araucaria muelleri in New Caledonia
Kinabalu, Borneo 53. Araucaria scopulorum seed cones in New Cal-
32. Agathis kinabaluensis sapling on the Mesilau edonia
River, Mt. Kinabalu, Borneo 54. Araucaria scopulorum foliage
33. Agathis lenticula tree at Kinabalu Park H. Q., 55. Athrotaxis cupressoides at Dove Lake, Tasma-
Borneo nia, Australia
34. Agathis lenticula trunk in Crocker Range, Bor- 56. Athrotaxis selaginoides at Dove Lake, Tasmania,
neo Australia
35. Agathis microstachya trees at Lake Barrine, 57. Athrotaxis selaginoides seed cones
Queensland, Australia 58. Austrocedrus chilensis forest on lava, Chile
36. Agathis ovata trees near Yaté, New Caledonia (photo M. Gardner)
37. Agathis ovata leaves and pollen cone 59. Austrocedrus chilensis seed cones

Second section (Fig. 60–120) 329–336

60. Austrotaxus spicata tree in New Caledonia 69. Callitris roei in Fitzgerald River N. P., Western
(photo M. Gardner) Australia
61. Callitris canescens in Western Australia 70. Callitris roei seed cones and foliage
62. Callitris columellaris in Kings Canyon N. P., 71. Calocedrus decurrens tree in the Sierra Nevada,
Northern Territories, Australia California, USA
63. Callitris macleayana trees in the Herberton 72. Calocedrus decurrens trunk in the Sierra Ne-
Range, Queensland, Australia vada, California, USA
64. Callitris macleayana seed cones 73. Calocedrus formosana foliage and seed cones
65. Callitris muelleri in New South Wales, Australia 74. Calocedrus macrolepis foliage and pollen cones
66. Callitris muelleri seed cones 75. Calocedrus rupestris foliage and pollen cones
67. Callitris preissii at Woodman Point, Western (photo L. Aveyanov)
Australia 76. Cathaya argyrophylla tree in Sichuan (photo
68. Callitris rhomboidea seed cones, Grampian H. Nimsch)
Mts., Victoria, Australia 77. Cathaya argyrophylla seed cones (photo
S. X. Yu)
78. Cedrus atlantica pollen cones 101. Dacrycarpus dacrydioides tree in North Island,
79. Cedrus deodara seed cones New Zealand
80. Cedrus libani var. libani in the Taurus Mts., 102. Dacrycarpus dacrydioides trunk
Turkey 103. Dacrycarpus dacrydioides seed cones
81. Cephalotaxus fortunei var. fortunei pollen cones 104. Dacrycarpus imbricatus var. imbricatus flushing
and foliage foliage
82. Cephalotaxus fortunei var. fortunei ripe seeds 105. Dacrycarpus kinabaluensis tree on Mt Kinaba-
83. Cephalotaxus harringtonii var. harringtonii pol- lu, Borneo
len cones 106. Dacrycarpus kinabaluensis seed cones
84. Cephalotaxus harringtonii var. harringtonii ripe 107. Dacrydium araucaroides in New Caledonia
seeds 108. Dacrydium araucarioides foliage and pollen 1067
85. Cephalotaxus mannii leaves and seeds (photo cones
L. Averyanov) 109. Dacrydium beccarii foliage with pollen cones
86. Chamaecyparis formosensis and C. obtusa in 110. Dacrydium comosum canopy at Gunung Ulu
Chilan Shan, Taiwan Kali, Malaysia
87. Chamaecyparis formosensis foliage and seed 111. Dacrydium comosum at Gunung Ulu Kali, Ma-
cones laysia
88. Chamaecyparis lawsoniana pollen cones 112. Dacrydium comosum foliage
89. Chamaecyparis lawsoniana seed cones 113. Dacrydium cupressinum in North Island, New
90. Chamaecyparis thyoides var. thyoides trunk in Zealand
North Carolina, USA 114. Dacrydium elatum at Gunung Ledang, Malay-
91. Cryptomeria japonica seed cones sia
92. Cunninghamia konishii foliage 115. Dacrydium gibbsiae small tree on Mt Kinabalu,
93. Cunninghamia lanceolata seed cones Borneo
94. Cupressus arizonica var. arizonica seed cones 116. Dacrydium gracile at Bukit Tupai, Mt Kinabalu,
95. Cupressus dupreziana tree in the Sahara (Tassili Borneo
n’Ajjer, Algeria) 117. Dacrydium guillauminii in New Caledonia
96. Cupressus guadalupensis var. forbesii in Califor- (photo A. Schmidt)
nia, USA 118. Dacrydium xanthandrum tree in the Crocker
97. Cupressus lusitanica var. benthamii seed cones Range, Borneo
98. Cupressus macrocarpa near Monterey, Califor- 119. Dacrydium xanthandrum trunk in the Crocker
nia, USA Range, Borneo
99. Cupressus torulosa var. torulosa seed cones 120. Diselma archeri in Cradle Mountain N. P., Tas-
100. Dacrycarpus cinctus foliage and cones (photo mania, Australia
T. Utteridge)

Third section (Fig. 121–176) 449–456

121. Falcatifolium falciforme trees in the Crocker 129. Halocarpus bidwillii in North Island, New Zea-
Range, Borneo land
122. Falcatifolium falciforme flushing leaves 130. Halocarpus bidwillii foliage
123. Falcatifolium falciforme seedling at Fraser’s Hill, 131. Halocarpus biformis foliage
Malaysia 132. Juniperus californica in Anza Borrego Desert
124. Falcatifolium taxoides on Mt. Panié, New Cal- State Park, California, USA
edonia 133. Juniperus californica seed cones
125. Fitzroya cupressoides in the N. P. Alerce Andi- 134. Juniperus chinensis var. sargentii foliage and
no, Chile (photo P. Woltz) seed cones
126. Fitzroya cupressoides foliage and seed cones 135. Juniperus communis var. communis foliage and
127. Fokienia hodginsii foliage and cones seed cones
128. Glyptostrobus pensilis pollen cones and seed 136. Juniperus communis var. saxatilis in Mt. Rainier
cones (photo D. White) N. P., Washington, USA
 137. Juniperus deppeana var. deppeana in Puebla, 155. Larix decidua var. decidua in the Alps, Switzer-
Mexico land
138. Juniperus deppeana var. deppeana trunk with 156. Larix decidua var. decidua bark
bark 157. Larix decidua var. decidua seed cones
139. Juniperus flaccida var. flaccida tree in Oaxaca, 158. Larix gmelinii var. principis-rupprechtii seed
Mexico cones
140. Juniperus occidentalis var. australis tree in the 159. Larix griffithii var. griffithii seed cone
Sierra Nevada, California, USA 160. Larix kaempferi seed cones
141. Juniperus oxycedrus subsp. macrocarpa foliage 161. Larix lyallii in the Wenatchee Mts., Washing-
and seed cones ton, USA
1068 142. Juniperus phoenicea subsp. phoenicea at Cape 162. Lepidothamnus fonkii in Chile (photo M. Gard-
St. Vincent, Portugal ner)
143. Juniperus phoenicea subsp. phoenicea foliage 163. Libocedrus bidwillii in North Island, New Zea-
and cones land
144. Juniperus pseudosabina in the Alaj Mountains, 164. Libocedrus bidwillii foliage and seed cones
Kirgyzstan 165. Libocedrus plumosa foliage
145. Juniperus pseudosabina seed cones in Kir- 166. Manoao colensoi seed cones (photo B. P. J. Mol-
gyzstan loy)
146. Juniperus sabina var. sabina in Kirgyzstan 167. Metasequoia glyptostroboides bark
147. Juniperus semiglobosa in Kirgyzstan 168. Metasequoia glyptostroboides seed cones
148. Juniperus semiglobosa foliage and seed cones 169. Microbiota decussata foliage
149. Juniperus semiglobosa pollen cones 170. Microcachrys tetragona foliage and seed cones
150. Juniperus virginiana var. virginiana tree in 171. Nageia fleuryi leaves and seed cones (photo
North Carolina, USA L. Averyanov)
151. Keteleeria davidiana var. davidiana foliage and 172. Nageia nagi flushing leaves
seed cones 173. Nageia wallichiana tree in Vietnam (photo
152. Keteleeria davidiana var. davidiana seed cone L. Averyanov)
153. Lagarostrobos franklinii tree at Riveaux Creek, 174. Nageia wallichiana leaves and seed cones (pho-
Tasmania, Australia to L. Averyanov)
154. Lagarostrobos franklinii foliage and pollen 175. Neocallitropsis pancheri in New Caledonia
cones 176. Neocallitropsis pancheri foliage

Fourth section (Fig. 177–239) 625–632

177. Nothotsuga longibracteata in Nan Ling Mts. 185. Phyllocladus hypophyllus canopy on Mt Kina-
Hunan, China balu, Borneo
178. Nothotsuga longibracteata seed cones (photo 186. Phyllocladus hypophyllus glaucous foliage on
Y. Liu) Mt. Kinabalu, Borneo at 3100 m
179. Papuacedrus papuana var. papuana pollen 187. Phyllocladus trichomanoides var. alpinus foliage
cones (photo D. White) 188. Picea chihuahuana green and ripe seed cones
180. Papuacedrus papuana leaves and seed cones 189. Picea likiangensis var. likiangensis young seed
(photo D. White) cones
181. Parasitaxus usta in New Caledonia (photo 190. Picea likiangensis var. likiangensis seed cones
W. Baker) (photo P. de Spoelberch)
182. Parasitaxus usta foliage and seed cones (photo 191. Picea likiangensis var. likiangensis seed cones
W. Baker) 192. Picea morrisonicola tree in Taroko N. P., Taiwan
183. Pherosphaera hookeriana in Tasmania, Austral- 193. Picea morrisonicola trunk with bark
ia 194. Picea orientalis foliage and pollen cones
184. Phyllocladus aspleniifolius foliage and pollen 195. Picea orientalis seed cones
cones 196. Picea schrenkiana subsp. tianschanica in Kir-
gyzstan
197. Picea schrenkiana subsp. tianschanica tree in 218. Pinus contorta var. murrayana in the Sierra Ne-
Kirgyzstan vada, California, USA
198. Picea sitchensis tree in Olympic N. P., Washing- 219. Pinus coulteri pollen cones
ton, USA 220. Pinus coulteri foliage and seed cones
199. Picea smithiana pollen cones 221. Pinus culminicola foliage
200. Picea smithiana seed cone from bud 222. Pinus devoniana foliage
201. Picea smithiana seed cone 223. Pinus durangensis in Durango, Mexico (photo
202. Picea wilsonii pollen cones C. Hughes)
203. Pilgerodendron uviferum pollen cones 224. Pinus engelmannii foliage
204. Pilgerodendron uviferum foliage and seed cones 225. Pinus hartwegii on Pico de Orizaba, Veracruz,
205. Pinus albicaulis foliage and pollen cones Mexico 1069
206. Pinus aristata foliage 226. Pinus heldreichii seed cones
207. Pinus armandii var. armandii seed cone 227. Pinus latteri forest in Thailand (photo H. Haze-
208. Pinus attenuata seed cones broek)
209. Pinus ayacahuite var. veitchii foliage and seed 228. Pinus longaeva on Telescope Peak, Death Valley
cones N. P., California, USA
210. Pinus balfouriana in the Sierra Nevada, Califor- 229. Pinus maximartinezii in Zacatecas, Mexico
nia, USA 230. Pinus maximartinezii seed cone and seedling
211. Pinus bungeana trunk with bark 231. Pinus monophylla foliage and seed cones
212 Pinus bungeana seed cone 232. Pinus monophylla leaves
213. Pinus cembra in the Alps, Switzerland 233. Pinus muricata pollen cones
214. Pinus cembra foliage and seed cones 234. Pinus muricata seed cones
215. Pinus cembroides var. cembroides in Hidalgo, 235. Pinus nelsonii seed cone
Mexico 236. Pinus nigra subsp. laricio trunk with bark
216. Pinus cembroides var. cembroides bark 237. Pinus nigra subsp. salzmannii seed cones
217. Pinus cembroides subsp. orizabensis seed cones 238. Pinus patula var. patula tree in Oaxaca, Mexico
239. Pinus patula var. patula leaves

Fifth section (Fig. 240–303) 785–792

240. Pinus pinaster subsp. pinaster pollen cones 257. Podocarpus brevifolius on the Mesilau River,
241. Pinus pinaster subsp. pinaster seed cones of two Mt. Kinabalu, Borneo
ages 258. Podocarpus brevifolius leaves
242. Pinus pinea in Algarve, Portugal 259. Podocarpus chingianus foliage and seed cone
243. Pinus pinea seed cone 260. Podocarpus costalis foliage
244. Pinus ponderosa var. ponderosa pollen cones 261. Podocarpus cunninghamiii foliage and seed
245. Pinus pungens seed cones cones
246. Pinus quadrifolia in California, USA 262. Podocarpus dispermus leaves
247. Pinus rzedowskii tree in Michoacán, Mexico 263. Podocarpus elatus foliage
248. Pinus rzedowskii seed cones 264. Podocarpus elatus seed cone
249. Pinus sylvestris var. sylvestris trees 265. Podocarpus grayae small sapling at Cape Tribu-
250. Pinus taiwanensis var. taiwanensis in Hehuan lation, Queensland, Australia
Shan, Taiwan 266. Podocarpus grayae tree at Lake Eacham,
251. Pinus virginiana seed cones Queensland, Australia
252. Pinus wallichiana var. wallichiana seed cones 267. Podocarpus grayae trunk, Herberton Range,
253. Platycladus orientalis foliage and seed cones Queensland, Australia
(photo D. Mabberley) 268. Podocarpus henkelii leaves
254. Platycladus orientalis seed cones 269. Podocarpus laubenfelsii seedling on Mt. Kina-
255. Podocarpus acutifolius foliage and pollen cones balu, Borneo
256. Podocarpus brassii var. brassii seed cone (photo 270. Podocarpus latifolius in the Drakensberg, South
T. Waters) Africa (photo J. Grimshaw)
 271. Podocarpus lawrencei seed cones 288. Podocarpus salignus foliage and young seed
272. Podocarpus macrophyllus var. macrophyllus fo- cones
liage 289. Podocarpus spinulosus on North Stradbroke Is-
273. Podocarpus macrophyllus var. macrophyllus fo- land, Queensland, Australia
liage and pollen cones 290. Podocarpus spinulosus unripe seed cone (photo
274. Podocarpus matudae leaves and young seed G. Garruthers
cone 291. Podocarpus spinulosus ripe seed cones (photo
275. Podocarpus milanjianus on Mt. Elgon, Uganda G. Garruthers)
(photo D. L. Roberts) 292. Podocarpus sprucei foliage buds (photo P. Cribb
276. Podocarpus nakaii flushing leaves 1425)
1070 277. Podocarpus nakaii foliage and seed cones 293. Podocarpus totara tree in North Island, New
278. Podocarpus neriifolius var. neriifolius in Papua Zealand
New Guinea (photo T. Utteridge) 294. Podocarpus totara bark
279. Podocarpus neriifolius var. neriifolius seed cones 295. Podocarpus totara foliage and pollen cones
(photo T. Utteridge) 296. Prumnopitys andina foliage and pollen cones
280. Podocarpus nivalis in North Island, New Zea- 297. Prumnopitys andina seed cones
land 298. Prumnopitys ferruginoides foliage on Mt. Mou,
281. Podocarpus nivalis foliage and seed cones New Caledonia
282. Podocarpus novae-caledoniae in New Caledo- 299. Prumnopitys ladei trunk on Mt. Lewis, Queens-
nia land, Australia
283. Podocarpus novae-caledoniae foliage and young 300. Prumnopitys ladei foliage
seed cones 301. Pseudolarix amabilis pollen cones
284. Podocarpus nubigenus leaves (upperside) 302. Pseudolarix amabilis seed cones
285. Podocarpus nubigenus leaves (underside) 303. Pseudotaxus chienii foliage and buds of seed
286. Podocarpus polystachyus pollen cones cones
287. Podocarpus rumphii seed cones

Sixth section (Fig. 304–362) 961–968

304. Pseudotsuga japonica foliage and young seed 316. Saxegothaea conspicua foliage with pollen
cone cones and seed cones
305. Pseudotsuga japonica seed cones 317. Sciadopitys verticillata foliage and pollen cone
306. Pseudotsuga menziesii tree in Mt. Rainier N. P., buds
Washington, USA 318. Sciadopitys verticillata seed cone (photo
307. Pseudotsuga menziesii giant trunk in Olympic C. N. Page)
N. P., Washington, USA 319. Sequoia sempervirens at Bull Creek, California,
308. Pseudotsuga menziesii var. menziesii seed cones USA (photo R. Van Pelt)
309. Pseudotsuga sinensis var. sinensis trees in Tar- 320. Sequoia sempervirens seed cones
oko N. P., Taiwan 321. Sequoiadendron giganteum in California, USA
310. Pseudotsuga sinensis var. sinensis fallen seed (photo E. Parker)
cones 322. Sequoiadendron giganteum pollen cones and
311. Retrophyllum comptonii young tree on Mt. seed cones
Panié, New Caledonia 323. Sundacarpus amarus trunk and bark, Lake Bar-
312. Retrophyllum comptonii foliage and seed (photo rine, Queensland, Australia
M. Gardner) 324. Sundacarpus amarus foliage
313. Retrophyllum minus in Rivière des Lacs, New 325. Sundacarpus amarus pollen cones
Caledonia 326. Taiwania cryptomerioides in Yunnan, China
314. Retrophyllum minus foliage and immature (photo D. Long)
seeds 327. Taiwania cryptomerioides tree (photo P. Tho-
315. Saxegothaea conspicua tree (photo C. N. Page) mas)
328. Taiwania cryptomerioides trunk in Taiwan
329. Taxodium distichum swamp forest in North 346. Thujopsis dolabrata var. dolabrata foliage (un-
Carolina, USA derside)
330. Taxodium distichum var. distichum foliage and 347. Torreya californica foliage with pollen cones
seed cones 348. Torreya californica foliage with seeds
331. Taxodium mucronatum in Oaxaca, Mexico 349. Torreya nucifera foliage with seeds
332. Taxus baccata ancient tree on ruined wall of 350. Tsuga chinensis var. chinensis in the mountains
Waverley Abbey, England of Taiwan
333. Taxus baccata foliage and pollen cones 351. Tsuga chinensis var. chinensis foliage and seed
334. Taxus baccata foliage and seeds cones
335. Taxus brevifolia in the Wenatchee Mts., Wash- 352. Tsuga forrestii foliage and seed cones
ington, USA 353. Tsuga heterophylla on a ‘nurse log’ in Olympic 1071
336. Taxus brevifolia foliage and seeds N. P., Washington, USA
337. Taxus cuspidata var. cuspidata in Korea (photo 354. Tsuga mertensiana var. mertensiana foliage and
Y. S. Kim) seed cones
338. Taxus cuspidata var. cuspidata foliage and seeds 355. Widdringtonia cedarbergensis seed cones
339. Tetraclinis articulata seed cones (photo 356. Widdringtonia nodiflora seed cones
M. Gardner) 357. Widdringtonia whytei on Mt. Mulanje, Malawi
340. Thuja koraiensis in Korea (photo Y. S. Kim) (photo P. Cribb)
341. Thuja koraiensis foliage branch (upperside) 358. Wollemia nobilis trees in Wollemi N. P., New
342. Thuja koraiensis foliage branch (underside) South Wales, Australia (photo J. Plaza)
343. Thuja plicata foliage and seed cones 359. Wollemia nobilis bud and foliage
344. Thuja sutchuenensis in Daba Shan, China (© 360. Wollemia nobilis foliage and pollen cones
Bedgebury Pinetum) 361. Xanthocyparis nootkatensis trunk in Olympic
345. Thujopsis dolabrata var. dolabrata foliage (up- N. P., Washington, USA
perside) 362. Xanthocyparis vietnamensis foliage
INDEX TO BOTANICAL NAMES OF CONIFERS

Names are listed at the ranks of family, genus, spe-

cies, subspecies, variety and forma. Names in bold
italics are accepted names with a descriptive ac-
count in this handbook, other names are synonyms 1073
or names of taxa that occur only in cultivation. Page
numbers are in bold where an account of the taxon
begins, underlined where an illustration is given.

Abies balsamea (L.) Mill. f. phanerolepis (Fernald)

A
Rehd. 63
Abies Mill. 17, 25, 38, 49, 50, 55, 56, 77, 90, 97, 109, Abies balsamea (L.) Mill. subsp. phanerolepis (Fern-
115, 118, 127, 262, 263, 266, 308, 403, 404, 422, ald) E. Murray 63
423, 436, 444, 445, 459, 463, 495, 554, 574, 578, Abies balsamea (L.) Mill. var. phanerolepis 63
592, 633, 666, 668, 754, 761, 796, 932, 933, 981, Abies beshanzuensis M. H. Wu 56, 58, 63, 128,
984, 1014, 1015 1063
Abies alba Mill. 55, 56, 57, 59, 60, 65, 68, 69, 103, Abies beshanzuensis M. H. Wu var. ziyuanensis
112, 177, 555, 591, 732 (L. K. Fu & S. L. Mo) L. K. Fu & Nan Li 127
Abies alba Mill. var. acutifolia Turrill 64, 68 Abies bicolor Maxim. 557
Abies alba Mill. subsp. apennina Brullo, Scelsi & Abies bifolia A. Murray bis var. arizonica (Merriam)
Spampinato 59 O’Kane & K. D. Heil 98
Abies alba Mill. subsp. borisii-regis (Mattf.) Kozuha- Abies ×borisii-regis Mattf. 56, 57, 64, 65, 68, 69,
rov & N. Andreev 64 732
Abies alba Mill. var. cephalonica (Loudon) Abies borisii-regis Mattf. var. pungenti-pilosa Viguié
Richt. 68 & Gaussen 64
Abies alba Mill. subsp. equi-trojani (Asch. & Sint. ex Abies bornmuelleriana Mattf. 107, 108
Boiss.) Asch. & Graebn. 108 Abies brachyphylla Maxim. 94
Abies alba Mill. subsp. nebrodensis (Lojac.) Abies brachyphylla Maxim. var. umbellata (Mayr)
Nitz. 102 Dallim. & A. B. Jacks. 94
Abies alba Mill. var. nebrodensis (Lojac.) Svobo- Abies brachytyla Franch. 563
da 102 Abies bracteata (D. Don) A. Poit. 56, 66, 67, 177
Abies alcoquiana Veitch ex Lindl. 557 Abies cephalonica Loudon 56, 57, 65, 68, 69, 112,
Abies alpestris Brügger 556 177, 409, 412, 416
Abies amabilis Douglas ex J. Forbes 56, 57, 61, 88, Abies chayuensis W. C. Cheng & L. K. Fu 85
113, 177, 600, 993, 1017, 1029 Abies chengii Rushforth 84, 85
Abies arizonica Merriam 98 Abies chensiensis Tiegh. 56, 58, 69, 70, 71, 82, 128,
Abies balsamea (L.) Mill. 56, 57, 62, 63, 401, 402, 562, 596
490, 582, 597, 991, 1009 Abies chensiensis Tiegh. subsp. chensiensis 70
Abies balsamea (L.) Mill. var. balsamea 63 Abies chensiensis Tiegh. var. ernestii (Rehd.)
Abies balsamea (L.) Mill. subsp. fraseri (Pursh) T. S. Liu 115
E. Murray 86 Abies chensiensis Tiegh. subsp. salouenensis (Bor-
Abies balsamea (L.) Mill. subsp. lasiocarpa (Hook.) dères & Gaussen) Rushforth 70, 84
Boivin 98 Abies chensiensis Tiegh. var. salouenensis (Bordères
Abies balsamea (L.) Mill. subsp. lasiocarpa (Hook.) & Gaussen) Silba 70
Boivin var. arizonica (Merriam) Boivin 98
 Abies chensiensis Tiegh. subsp. yulongxueshanensis Abies diversifolia Maxim. 1012
Rushforth 71 Abies durangensis Martínez 56, 58, 77, 78, 126,
Abies chensiensis Tiegh. var. yulongxueshanensis 566
(Rushforth) Silba 71 Abies durangensis Martínez var. coahuilensis
Abies chinensis Franch. 1011 (I. M. Johnst.) Martínez 78
Abies cilicica (Antoine & Kotschy) Carrière 56, 57, Abies durangensis Martínez var. durangensis 78
71, 72, 259, 409, 412, 416 Abies equi-trojani (Asch. & Sint. ex Boiss.) Mattf.
Abies cilicica (Antoine & Kotschy) Carrière var. 107, 108
borisii-regis (Mattf.) Silba 64 Abies ernestii Rehd. 70, 115
Abies cilicica (Antoine & Kotschy) Carrière subsp. Abies ernestii Rehd. var. salouenensis (Bordères &
1074 cilicica 71, 72 Gaussen) W. C. Cheng & L. K. Fu 70, 71
Abies cilicica (Antoine & Kotschy) Carrière subsp. Abies excelsior Franco 87
isaurica Coode & Cullen 71, 72 Abies fabri (Mast.) Craib 56, 59, 64, 79, 115
Abies cilicica (Antoine & Kotschy) Carrière var. Abies fabri (Mast.) Craib var. beshanzuensis (M. H.
pyramidalis Boydak & Erdogrul 72 Wu) Silba 63
Abies coahuilensis I. M. Johnst. 78 Abies fabri (Mast.) Craib subsp. fabri 79
Abies colimensis Rushforth & Narave 116 Abies fabri (Mast.) Craib subsp. minensis (Bordères
Abies concolor (Gordon) Lindl. ex Hildebr. 56, 58, & Gaussen) Rushforth 80
72, 73, 99, 247, 276, 292, 295, 564, 690, 715, 931, Abies fabri (Mast.) Craib var. ziyuanensis (L. K. Fu
951 & S. L. Mo) Silba 127
Abies concolor (Gordon) Lindl. ex Hildebr. f. atro­ Abies fanjingshanensis W. L. Huang 56, 59, 80
violacea Cinovskis 73 Abies fansipanensis Q. P. Xiang 75
Abies concolor (Gordon) Lindl. ex Hildebr. var. Abies fargesii Franch. 56, 59, 81, 82, 589, 595
baja­californica Silba 73 Abies fargesii Franch. var. fanjingshanensis (W. L.
Abies concolor (Gordon) Lindl. ex Hildebr. subsp. Huang) Silba 80
lowiana (Gordon) E. Murray 72 Abies fargesii Franch. var. fargesii 82
Abies concolor (Gordon) Lindl. ex Hildebr. var. Abies fargesii Franch. var. faxoniana (Rehd. &
lowiana (Gordon) Lemmon 72 E. H. Wilson) T. S. Liu 80, 82, 115, 122
Abies concolor (Gordon) Lindl. ex Hildebr. var. Abies fargesii Franch. var. hupehensis Silba 82
martinezii Silba 73 Abies fargesii Franch. var. sutchuenensis Franch.
Abies dayuanensis Q. X. Liu 127 70, 82
Abies delavayi Franch. 56, 59, 74, 75, 77, 85, 445, Abies faxoniana Rehd. & E. H. Wilson 82
1055 Abies ferreana Bordères & Gaussen 85
Abies delavayi Franch. var. delavayi 75, 177 Abies ferreana Bordères & Gaussen var. longibracte­
Abies delavayi Franch. var. fabri (Mast.) ata L. K. Fu & Nan Li 85
D. R. Hunt 79 Abies firma Siebold & Zucc. 56, 58, 64, 83, 94,
Abies delavayi Franch. subsp. fansipanensis 280, 285, 930, 947, 1005, 1019
(Q. P. Xiang) Rushforth 75 Abies firma Siebold & Zucc. var. brachyphylla
Abies delavayi Franch. var. faxoniana (Rehd. & (­Maxim.) Bertrand 94
E. H. Wilson) A. B. Jacks. 82 Abies flinckii Rushforth 90
Abies delavayi Franch. var. forrestii (Coltm.-Rog.) Abies fordei Rushforth 76, 77, 487
A. B. Jacks. 85 Abies forrestii Coltm.-Rog. 56, 75, 84, 85, 126, 127,
Abies delavayi Franch. var. georgei (Orr) Melville   85 445, 563, 955
Abies delavayi Franch. var. motuoensis W. C. Cheng Abies forrestii Coltm.-Rog. var. chayuensis (W. C.
& L. K. Fu 76 Cheng & L. K. Fu) Silba 85
Abies delavayi Franch. var. nukiangensis (W. C. Abies forrestii Coltm.-Rog. var. chengii (Rushforth)
Cheng & L. K. Fu) Farjon & Silba 76, 177 Silba 85
Abies delavayi Franch. var. smithii (Viguié & Gaus- Abies forrestii Coltm.-Rog. var. ferreana (Bordères
sen) T. S. Liu 86 & Gaussen) Farjon & Silba 85
Abies densa Griff. 56, 59, 76, 77, 445, 487, 563, 602, Abies forrestii Coltm.-Rog. var. forrestii 85
780
Abies forrestii Coltm.-Rog. var. georgei (Orr) Abies kawakamii (Hayata) T. Itô 55, 56, 58, 94, 95,
­Farjon 85 178, 587
Abies forrestii Coltm.-Rog. var. smithii Viguié & Abies koreana E. H. Wilson 56, 58, 95, 96, 178
Gaussen 86 Abies koreana E. H. Wilson f. nigrocarpa Hatus. 95
Abies fraseri (Pursh) Poir. 56, 58, 63, 86, 87, 597 Abies lasiocarpa (Hook.) Nutt. 56, 57, 61, 97, 99,
Abies gamblei Hickel 111 113, 401, 461, 491, 493, 568, 582, 600, 619, 654,
Abies georgei Orr 85 676, 715, 932, 993, 1017, 1029
Abies georgei Orr var. smithii (Viguié & Gaussen) Abies lasiocarpa (Hook.) Nutt. subsp. arizonica
W. C. Cheng & L. K. Fu 86 (Merriam) E. Murray 98
Abies glehnii F. Schmidt 572 Abies lasiocarpa (Hook.) Nutt. var. arizonica (Mer-
Abies gmelinii Rupr. 485 riam) Lemmon 97, 98 1075
Abies gracilis Kom. 118 Abies lasiocarpa (Hook.) Nutt. var. fallax (Engelm.)
Abies grandis (Douglas ex D. Don) Lindl. 56, 58, Franco 98
61, 73, 87, 113, 247, 277, 493, 698, 715, 949, 993 Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa 98
Abies grandis (Douglas ex D. Don) Lindl. var. ida­ Abies likiangensis Franch. 578
hoensis Silba 87 Abies lowiana (Gordon) A. Murray bis 72
Abies grandis (Douglas ex D. Don) Lindl. var. Abies lowiana (Gordon) A. Murray bis var. viridula
lowiana (Gordon) Hoopes 72 Debreczy & Rácz 73
Abies griffithiana Lindl. & Gordon 487 Abies macrocarpa Vasey 931
Abies guatemalensis Rehd. 56, 58, 88, 89, 125, 683, Abies magnifica A. Murray bis 56, 58, 61, 73, 98,
713, 771, 981 99, 100, 178, 247, 428, 564, 690, 698, 715, 951,
Abies guatemalensis Rehd. var. guatemalensis 1030
89, 90 Abies magnifica A. Murray bis var. magnifica
Abies guatemalensis Rehd. var. jaliscana Mar- 99, 100
tínez 90 Abies magnifica A. Murray bis var. shastensis
Abies guatemalensis Rehd. var. longibracteata De- ­Lemmon 99
breczy & Rácz 89 Abies mariana Mill. 581
Abies guatemalensis Rehd. var. tacanensis (Lundell) Abies mariesii Mast. 56, 57, 100, 101, 123, 557, 996,
Martínez 89 1013
Abies heterophylla Raf. 1015 Abies mariesii Mast. var. kawakamii Hayata 94
Abies hickelii Flous & Gaussen 56, 59, 89, 90, 92 Abies marocana Trab. 112
Abies hickelii Flous & Gaussen var. hickelii 91 Abies mayriana (Miyabe & Kudô) Miyabe &
Abies hickelii Flous & Gaussen var. macrocarpa Kudô 118
Martínez 91 Abies menziesii Mirb. 929, 933
Abies hickelii Flous & Gaussen var. oaxacana (Mar- Abies mexicana Martínez 126
tínez) Farjon & Silba 91 Abies minensis Bordères & Gaussen 80
Abies hidalgensis Debreczy, Rácz & Guízar 56, 59, Abies nebrodensis (Lojac.) Mattei 56, 57, 102, 103,
91, 92 178
Abies holophylla Maxim. 56, 58, 92, 93, 484, 693 Abies nemorensis (Mayr) Miyabe & Kudô 119
Abies holophylla Maxim. var. aspericorticea Y. Y. Abies neodurangensis Debreczy, Rácz & Salazar 78
Sun 92 Abies nephrolepis (Trautv. ex Maxim.) Maxim. 56,
Abies homolepis Siebold & Zucc. 56, 58, 83, 93, 94, 58, 93, 103, 104, 105, 484, 515, 560, 576, 586, 589,
128, 178, 489, 604, 994, 996, 1013 978, 990
Abies homolepis Siebold & Zucc. var. homo­lepis  94 Abies nephrolepis (Trautv. ex Maxim.) Maxim.
Abies homolepis Siebold & Zucc. var. umbellata subsp. sachalinensis (F. Schmidt) V. N. Voro-
(Mayr) E. H. Wilson 94 shilov 118
Abies intermedia Fulling 63 Abies nordmanniana (Steven) Spach 56, 57, 69,
Abies jezoensis Siebold & Zucc. 574 105, 106, 107, 112, 412, 416, 592
Abies kaempferi Lindl. 924 Abies nordmanniana (Steven) Spach subsp. born­
Abies kansouensis Bordères & Gaussen 82 muelleriana (Mattf.) Coode & Cullen 108
 Abies nordmanniana (Steven) Spach var. born­ Abies religiosa (Kunth) Schltdl. & Cham. 56, 59,
muelleriana (Mattf.) Silba 108 89, 90, 116, 125, 710, 713, 731, 1061
Abies nordmanniana (Steven) Spach subsp. equi- Abies religiosa (Kunth) Schltdl. & Cham. var. emar­
trojani (Asch. & Sint. ex Boiss.) Coode & ginata Loock ex Martínez 90
­Cullen 107, 108 Abies sachalinensis (F. Schmidt) Mast. 56, 58, 117,
Abies nordmanniana (Steven) Spach var. equi- 118, 266, 573, 978
trojani (Asch. & Sint. ex Boiss.) Guin. & Abies sachalinensis (F. Schmidt) Mast. f. corticosa
Maire 108 (Tatew.) Hayashi 118
Abies nordmanniana (Steven) Spach subsp. nord­ Abies sachalinensis (F. Schmidt) Mast. var. corticosa
manniana 108 Tatew. 118
1076 Abies nukiangensis W. C. Cheng & L. K. Fu 76 Abies sachalinensis (F. Schmidt) Mast. var. gracilis
Abies numidica de Lannoy ex Carrière 56, 59, (Kom.) Farjon 118
108, 109, 112, 178, 255 Abies sachalinensis (F. Schmidt) Mast. var. mayri­
Abies oaxacana Martínez 91 ana Miyabe & Kudô 118
Abies pectinata Guss. 102 Abies sachalinensis (F. Schmidt) Mast. var. nemo­
Abies pectinata Gilib. var. equi-trojani Asch. & Sint. rensis Mayr 119
ex Boiss. 108 Abies sachalinensis (F. Schmidt) Mast. var. sacha­
Abies pectinata Gilib. var. nebrodensis Lojac. 102 linensis 118, 484
Abies ×phanerolepis (Fernald) T. S. Liu 63 Abies salouenensis Bordères & Gaussen 70
Abies pindrow (Royle ex D. Don) Royle 56, 58, Abies semenovii B. Fedtsch. 120
109, 110, 112, 121, 258, 414, 462, 601 Abies shastensis (Lemmon) Lemmon 99
Abies pindrow (Royle ex D. Don) Royle var. brevi­ Abies sibirica Ledeb. 56, 57, 105, 119, 403, 484, 497,
folia Dallim. & A. B. Jacks. 111 590, 598, 759
Abies pindrow (Royle ex D. Don) Royle subsp. gam­ Abies sibirica Ledeb. var. gracilis (Kom.) Patschke   118
blei (Hickel) Rushforth 111 Abies sibirica Ledeb. var. nephrolepis Trautv. ex
Abies pindrow (Royle ex D. Don) Royle var. inter­ Maxim. 103
media 110 Abies sibirica Ledeb. subsp. semenovii (B. Fedtsch.)
Abies pindrow (Royle ex D. Don) Royle var. pin­ Farjon 120, 598
drow 110 Abies sibirica Ledeb. var. semenovii (B. Fedtsch.)
Abies pinsapo Boiss. 56, 59, 69, 111, 112, 179 T. S. Liu 120
Abies pinsapo Boiss. var. marocana (Trab.) Cebal- Abies sibirica Ledeb. subsp. sibirica 120
los & Bolaño 112, 255, 734 Abies ×sibiriconephrolepis Taken. & J. J. Chien
Abies pinsapo Boiss. subsp. numidica (de Lannoy ex 103, 105
Carrière) E. Murray 108 Abies sikokiana Nakai 124
Abies pinsapo Boiss. var. numidica (de Lannoy ex Abies spectabilis (D. Don) Mirb. 56, 59, 77, 110,
Carrière) Salomon 108 120, 121, 179, 258, 487, 601, 780
Abies pinsapo Boiss. var. pinsapo 112 Abies spectabilis (D. Don) Spach var. brevifolia
Abies pinsapo Boiss. subsp. tazaotana (S. Côzar ex (A. Henry) Rehd. 120
Villar) R. Govaerts 112 Abies spectabilis (D. Don) Spach var. densa (Griff.)
Abies pinsapo Boiss. var. tazaotana (S. Côzar ex Silba 76
­Villar) Pourtet 112 Abies spectabilis (D. Don) Spach var. langtangensis
Abies polita Siebold & Zucc. 602 Silba 120
Abies procera Rehd. 56, 58, 73, 99, 100, 113, 179, Abies spinulosa Griff. 601
698, 715, 993, 1029 Abies squamata Mast. 56, 57, 114, 122, 179, 562, 596
Abies recurvata Mast. 56, 58, 70, 82, 114, 115, 122, Abies subalpina Engelm. var. fallax Engelm. 98
179, 562 Abies sutchuenensis (Franch.) Rehd. & E. H. Wil-
Abies recurvata Mast. var. ernestii (Rehd.) C. T. son 82
Kuan 115 Abies tacanensis Lundell 89, 125
Abies recurvata Mast. var. recurvata 115 Abies tazaotana S. Côzar ex Villar 112
Abies recurvata Mast. var. salouenensis (Bordères & Abies torano Siebold ex K. Koch 602
Gaussen) C. T. Kuan 70 Abies tsuga Siebold & Zucc. 1007
Abies umbellata Mayr 94 Afrocarpus dawei (Stapf) C. N. Page 137
Abies veitchii Lindl. 56, 58, 94, 100, 123, 489, 557, Afrocarpus falcatus (Thunb.) C. N. Page 136, 137,
1013 138, 139, 140, 180, 844, 850
Abies veitchii Lindl. var. komagatakensis Hayashi   124 Afrocarpus gaussenii (Woltz) C. N. Page 138
Abies veitchii Lindl. f. olivacea (Shiras.) Cinovs­ Afrocarpus gracilior (Pilg.) C. N. Page 137, 139,
kis 124 140, 180, 439
Abies veitchii Lindl. var. olivacea Shiras. 124 Afrocarpus mannii (Hook. f.) C. N. Page 141
Abies veitchii Lindl. var. sachalinensis F. Schmidt 118 Afrocarpus usambarensis (Pilg.) C. N. Page 137,
Abies veitchii Lindl. var. sikokiana (Nakai) Ku- 141, 142, 844
saka 124 Agathis Salisb. 19, 26, 29, 33, 44, 143, 144, 148, 151,
Abies veitchii Lindl. var. veitchii 124 153, 157, 158, 163, 167, 321, 324, 343, 347, 366, 507, 1077
Abies vejarii Martínez 56, 59, 124, 125, 126, 566, 524, 540, 546, 812, 864, 868, 907, 938, 953, 1026
583, 680 Agathis atropurpurea B. Hyland 144, 145, 146, 159,
Abies vejarii Martínez var. macrocarpa Martínez 160
125 Agathis australis (D. Don) Lindl. 19, 143, 144, 146,
Abies vejarii Martínez subsp. mexicana (Martínez) 147, 158, 180, 322, 346, 380, 507, 548, 549, 827,
Farjon 125, 126 908, 916
Abies vejarii Martínez var. mexicana (Martínez) Agathis borneensis Warb. 145, 148, 149, 150, 151,
T. S. Liu 57, 125, 126 152, 153, 157, 163, 180, 350, 365, 851
Abies vejarii Martínez var. vejarii 125 Agathis brownii (Lem.) L. H. Bailey 157
Abies ×vilmorinii Matf. 112 Agathis celebica (Koord.) Warb. 150, 151
Abies webbiana (Wall. ex D. Don) Lindl. var. brevi­ Agathis celebica (Koord.) Warb. subsp. flavescens
folia A. Henry 120 (Ridl.) Veldkamp & Whitmore 152
Abies webbiana (Wall. ex D. Don) Lindl. var. pin­ Agathis corbassonii de Laub. 161, 162
drow (Royle ex D. Don) Brandis 110 Agathis dammara (Lamb.) Rich. & A. Rich. 143,
Abies yuanbaoshanensis Y. J. Lu & L. K. Fu 56, 59, 145, 148, 150, 151, 152, 153, 157, 366
126, 128 Agathis dammara (Lamb.) Rich. & A. Rich. subsp.
Abies yunnanensis Franch. 1013 dammara 154
Abies zapotekensis Debreczy 125 Agathis dammara (Lamb.) Rich. & A. Rich. subsp.
Abies zapotekensis Debreczy, Rácz & Ramírez 89 flavescens (Ridl.) Whitmore 152
Abies ziyuanensis L. K. Fu & S. L. Mo 56, 58, 127, Agathis endertii Meijer Drees 148
128 Agathis flavescens Ridl. 144, 152, 153, 154
Abietia A. H. Kent 929 Agathis hypoleuca (C. Moore ex Henkel &
Abietia douglasii (Sabine ex D. Don) A. H. Kent 929 W. Hochst.) Warb. 163
Acmopyle Pilg. 40, 51, 52, 129, 130, 337 Agathis kinabaluensis de Laub. 145, 153, 154, 163,
Acmopyle alba J. T. Buchholz 129 181
Acmopyle pancheri (Brongn. & Gris) Pilg. 129, Agathis labillardierei Warb. 145, 154, 155, 810, 943
130, 179, 938 Agathis lanceolata (Sébert & Pancher) Warb. 144,
Acmopyle sahniana J. T. Buchholz & N. E. Gray 155, 156, 196, 214, 328
129, 130, 131 Agathis lenticula de Laub. 145, 156, 157, 163, 181
Actinostrobus Miq. 36, 46, 48, 132, 135 Agathis macrophylla (Lindl.) Mast. 144, 157, 158,
Actinostrobus acuminatus Parl. 132, 134 167, 943
Actinostrobus arenarius C. A. Gardner 132, 133, Agathis macrophylla (Lindl.) Mast. var. obtusa
134, 135, 179, 180 (Lindl.) Silba 157
Actinostrobus pyramidalis Miq. 132, 134, 135 Agathis microstachya J. F. Bailey &
Actinostrobus pyramidalis Miq. subsp. arenarius C. T. White 144, 159, 160, 181
(C. A. Gardner) Silba 133 Agathis montana de Laub. 144, 160, 161, 211, 369
Actinostrobus pyramidalis Miq. var. arenarius Agathis moorei (Lindl.) Mast. 144, 161, 162
(C. A. Gardner) Silba 133 Agathis obtusa (Lindl.) Mast. 157
Afrocarpus (J. T. Buchholz & N. E. Gray) C. N. Page Agathis orbicula de Laub. 144, 162, 163
18, 22, 26, 29, 39, 51, 52, 136, 140, 844
 Agathis ovata (C. Moore ex Vieill.) Warb. 130, Araucaria araucana (Molina) K. Koch 19, 27, 182,
144, 163, 164, 181, 210, 340, 341, 526, 871 185, 186, 187, 188, 191, 192, 193, 200
Agathis philippinensis Warb. 150, 151 Araucaria bernieri J. T. Buchholz 182, 186, 188,
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey 193, 194, 212
144, 145, 165 Araucaria bernieri J. T. Buchholz var. pumilio
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey Silba 211, 212
subsp. nesophila Whitmore 165, 166 Araucaria bidwillii Hook. 19, 27, 44, 182, 183, 185,
Agathis robusta (C. Moore ex F. Muell.) F. M. Bailey 186, 187, 191, 194, 195, 198
subsp. robusta 165, 166 Araucaria biramulata J. T. Buchholz 186, 187, 195
Agathis silbae de Laub. 144, 158, 166, 167 Araucaria columnaris (J. R. Forst.) Hook. 183,
1078 Agathis spathulata de Laub. 166 186, 188, 196, 197, 206, 209
Agathis vitiensis (Seem.) Benth. & Hook. f. 157 Araucaria columnaris (G. Forst.) Hook. f. luxurians
Amentotaxus Pilg. 20, 41, 45, 53, 168, 175, 270 (Brongn. & Gris) E. H. Wilson 205
Amentotaxus argotaenia (Hance) Pilg. 168, 169, Araucaria cookii R. Br. ex Lindl. var. luxurians
170, 172, 173, 176, 1032 Brongn. & Gris 205
Amentotaxus argotaenia (Hance) Pilg. var. argo­ Araucaria cunninghamii Aiton ex A. Cunn. 183,
taenia 169 186, 187, 195, 198, 199, 203, 815
Amentotaxus argotaenia (Hance) Pilg. var. brevi­ Araucaria cunninghamii Aiton ex A. Cunn. var.
folia K. M. Lan & F. H. Zhang 170 cunninghamii 199
Amentotaxus argotaenia (Hance) Pilg. var. cathay­ Araucaria cunninghamii Aiton ex A. Cunn. var.
ensis (H. L. Li) P. C. Keng 169 papuana Lauterb. 199, 537
Amentotaxus argotaenia (Hance) Pilg. var. yunnan­ Araucaria heterophylla (Salisb.) Franco 183, 186,
ensis (H. L. Li) P. C. Keng 175 188, 200, 201
Amentotaxus assamica D. K. Ferguson 168, 170, Araucaria humboldtensis J. T. Buchholz 186, 188,
171 201, 205, 352, 369, 504
Amentotaxus cathayensis H. L. Li 169 Araucaria hunsteinii K. Schum. 185, 186, 187, 198,
Amentotaxus formosana H. L. Li 168, 171, 172, 175, 199, 202, 203
182 Araucaria hunsteinii K. Schum. var. klinkii
Amentotaxus hatuyenensis Hiep 168, 173, 174, 175, (­Lauterb.) Silba 202
251 Araucaria klinkii Lauterb. 202, 203
Amentotaxus poilanei (Ferré & Rouane) D. K. Fer- Araucaria laubenfelsii Corbasson 183, 186, 187,
guson 168, 174, 175 202, 204, 352, 369
Amentotaxus yunnanensis H. L. Li 168, 169, 171, Araucaria luxurians (Brongn. & Gris) de Laub.
173, 174, 175, 176, 182, 251 186, 187, 205, 206
Amentotaxus yunnanensis H. L. Li var. formosana Araucaria montana Brongn. & Gris 186, 187, 202,
(H. L. Li) Silba 171 205, 206, 369
Amentotaxus yunnanensis H. L. Li var. poilanei Araucaria muelleri (Carrière) Brongn. & Gris
Ferré & Rouane 174 183, 186, 187, 207, 208, 340
Americus Hanford 950 Araucaria nemorosa de Laub. 186, 187, 197, 208,
Americus gigantea (Lindl.) Hanford 950 209
Apinus Neck. ex Rydb. 608 Araucaria rulei F. Muell. 186, 187, 209, 210
Apinus cembra Neck. ex Rydb. 608 Araucaria schmidii de Laub. 161, 186, 187, 210, 211
Araucaria Juss. 19, 22, 27, 33, 44, 130, 164, 185, 186, Araucaria schumanniana Warb. 202, 203
190, 191, 201, 207, 209, 211, 340, 341, 524, 526, Araucaria scopulorum de Laub. 184, 186, 187, 211,
882, 938, 1026, 1027 212, 213
Araucaria angustifolia (Bertol.) Kuntze 19, 186, Araucaria subulata Vieill. 156, 186, 187, 193, 214
187, 188, 189, 190, 192, 1056 Araucariaceae Henkel & W. Hochst. 15, 16, 19,
Araucaria angustifolia (Bertol.) Kuntze var. depend­ 22, 26, 33, 44, 143, 185, 204, 524, 916, 938, 1026,
ens J. R. de Mattos 188 1027, 1057, 1058, 1061
Araucaria angustifolia (Bertol.) Kuntze var. vinacea Arceuthos Antoine & Kotschy 381, 409
J. R. de Mattos 188
Arceuthos drupacea (Labill.) Antoine & Kotschy Callitris baileyi C. T. White 224, 225
381, 407 Callitris canescens (Parl.) S. T. Blake 224, 225,
Arthrotaxis Endl. 216 226, 329
Athrotaxis D. Don 34, 46, 47, 216 Callitris columellaris F. Muell. 224, 225, 227, 228,
Athrotaxis cupressoides D. Don 184, 216, 218, 362, 239, 329
516, 543, 545 Callitris columellaris F. Muell. var. campestris
Athrotaxis ×laxifolia Hook. 216, 217, 545 Silba 227
Athrotaxis selaginoides D. Don 184, 216, 217, 218, Callitris columellaris F. Muell. var. intratropica
219, 545 (R. T. Baker & H. G. Smith) Silba 227
Athrotaxis tetragona Hook. 516 Callitris columellaris F. Muell. var. microcarpa
Austrocedrus Florin & Boutelje 36, 46, 48, 220 (Benth.) Govaerts 227 1079
Austrocedrus chilensis (D. Don) Pic. Serm. & Biz- Callitris drummondii (Parl.) F. Muell. 224, 225,
zarri 184, 192, 220, 221, 914, 1057 229
Austrotaxus R. H. Compton 41, 53, 222 Callitris endlicheri (Parl.) F. M. Bailey 224, 225,
Austrotaxus spicata R. H. Compton 222, 329 228, 230, 239, 240
Callitris glaucophylla J. Thompson & L. A. S. John-
son 227, 228
B
Callitris gracilis R. T. Baker subsp. murrayensis
Biota (D. Don) Endl. 793 (J. Garden) K. D. Hill 238, 239, 298
Biota orientalis (L.) Endl. 793 Callitris intratropica R. T. Baker & H. G. Smith
Bracteocarpus A. V. Bobrov & Melikyan 317, 320 227, 228
Bracteocarpus cinctus (Pilg.) A. V. Bobrov & Callitris macleayana (F. Muell.) F. Muell. 224,
­Melikyan 318 231, 242, 329
Bracteocarpus compactus (Wasscher) A. V. Bobrov Callitris monticola J. Garden 224, 225, 232, 233
& Melikyan 319 Callitris muelleri (Parl.) Benth. & Hook. f. ex
Bracteocarpus cumingii (Parl.) A. V. Bobrov & F. Muell. 224, 225, 234, 329
­Melikyan 320 Callitris neocaledonica Dummer 202, 224, 225,
Bracteocarpus dacrydiifolius (Wasscher) A. V. Bo- 235, 526
brov & Melikyan 318 Callitris oblonga Rich. & A. Rich. 224, 225, 236,
Bracteocarpus expansus (de Laub.) A. V. Bobrov & 237, 1059
Melikyan 322 Callitris oblonga Rich. & A. Rich. subsp. corangensis
Bracteocarpus imbricatus (Blume) A. V. Bobrov & K. D. Hill 236
Melikyan 317, 324 Callitris oblonga Rich. & A. Rich. subsp. parva
Bracteocarpus kawaii (Hayata) A. V. Bobrov & K. D. Hill 236
­Melikyan 324 Callitris preissii Miq. 134, 135, 224, 225, 238, 239,
Bracteocarpus kinabaluensis (Wasscher) A. V. Bo- 240, 241, 243, 245, 330
brov & Melikyan 325 Callitris preissii Miq. subsp. murrayensis J. Gar-
Bracteocarpus leptophyllus (Wasscher) A. V. Bobrov den 238
& Melikyan 351 Callitris preissii Miq. var. murrayensis (J. Garden)
Bracteocarpus papuanus (Ridl.) A. V. Bobrov & Silba 238
­Melikyan 325 Callitris preissii Miq. subsp. verrucosa (A. Cunn. ex
Bracteocarpus steupii (Wasscher) A. V. Bobrov & Endl.) J. Garden 243
Melikyan 326 Callitris preissii Miq. var. verrucosa (A. Cunn. ex
Brownetera Rich. ex Tratt. 544 Endl.) Silba 243
Brownetera aspleniifolia (Labill.) Tratt. 544 Callitris rhomboidea R. Br. ex Rich. & A. Rich.
Brunia nodiflora L. 1020, 1022 224, 225, 230, 234, 239, 330
Callitris roei (Endl.) F. Muell. 134, 135, 224, 225,
240, 330
C
Callitris schwarzii Marloth 1023
Callitris Vent. 19, 37, 46, 48, 224, 232, 239, 242, Callitris sulcata (Parl.) Schltr. 224, 225, 241, 242
525, 1057
 Callitris tuberculata R. Br. ex R. T. Baker & H. G. Cedrus deodara (Lamb.) G. Don 110, 254, 256,
Smith 238, 243 257, 258, 331, 414, 462, 601, 754, 780, 984, 1014
Callitris verrucosa (A. Cunn. ex Endl.) F. ­Muell. Cedrus libani A. Rich. 72, 254, 255, 256, 258, 259,
224, 225, 239, 243, 244 260, 331, 409, 412, 416, 431
Callitropsis Oerst. 290, 291, 1028, 1029 Cedrus libani A. Rich. subsp. atlantica (Endl.) Batt.
Callitropsis R. H. Compton 525 & Trab. 254
Callitropsis abramsiana (C. B. Wolf) D. P. Little   305 Cedrus libani A. Rich. var. atlantica (Endl.) Hook.
Callitropsis araucarioides R. H. Compton 525 f. 254
Callitropsis arizonica (Greene) D. P. Little 293 Cedrus libani A. Rich. subsp. brevifolia (Hook. f.)
Callitropsis bakeri (Jeps.) D. P. Little 295 Meikle 260
1080 Callitropsis benthamii (Endl.) D. P. Little 308 Cedrus libani A. Rich. var. brevifolia Hook. f. 260
Callitropsis forbesii (Jeps.) D. P. Little 306 Cedrus libani A. Rich. subsp. deodara (Lamb.)
Callitropsis glabra (Sudw.) D. P. Little 293 P. D. Sell 256
Callitropsis goveniana (Gordon) D. P. Little 304 Cedrus libani A. Rich. var. libani 259, 260
Callitropsis guadalupensis (S. Watson) D. P. Lit- Cedrus libani A. Rich. subsp. stenocoma (O. Schwarz)
tle 306 P. H. Davis 259
Callitropsis lusitanica (Mill.) D. P. Little 308 Cedrus libani A. Rich. var. stenocoma (O. Schwarz)
Callitropsis macnabiana (Hartw.) D. P. Little 309 Frankis 259
Callitropsis macrocarpa (Hartw.) D. P. Little 311 Cedrus libanitica Trew ex Pilg. subsp. stenocoma
Callitropsis montana (Wiggins) D. P. Little 293 O. Schwarz 259
Callitropsis nevadensis (Abrams) D. P. Little 294 Cephalotaxaceae Neger 16, 19, 22, 24, 33, 43, 44,
Callitropsis nootkatensis (D. Don) D. P. Little 1028, 45, 251, 261, 1054, 1062
1029 Cephalotaxus Siebold & Zucc. ex Endl. 14, 33, 44,
Callitropsis pigmaea (Lemmon) D. P. Little 304 45, 169, 261, 265, 270, 495
Callitropsis sargentii (Jeps.) D. P. Little 312 Cephalotaxus alpina (H. L. Li) L. K. Fu 263
Callitropsis stephensonii (C. B. Wolf) D. P. Little  294 Cephalotaxus argotaenia (Hance) Pilg. 169
Callitropsis vietnamensis (Farjon & Hiep) D. P. Lit- Cephalotaxus celebica Warb. 983
tle 1030 Cephalotaxus drupacea Siebold & Zucc. 265, 266,
Calocedrus Kurz 35, 46, 47, 246, 250, 503 273
Calocedrus decurrens (Torr.) Florin 88, 99, 221, Cephalotaxus drupacea Siebold & Zucc. var. har­
246, 247, 250, 292, 295, 330, 428, 564, 690, 698, ringtonii (Knight ex J. Forbes) Pilg. 266
715, 951, 993, 1030 Cephalotaxus drupacea Siebold & Zucc. var. sinensis
Calocedrus formosana (Florin) Florin 246, 248, Rehd. & E. H. Wilson 272
250, 275, 280, 287, 330, 955 Cephalotaxus fortunei Hook. 261, 262, 268, 269,
Calocedrus macrolepis Kurz 246, 248, 249, 250, 289, 331, 374, 475, 476, 522, 858, 955, 1003, 1015
330 Cephalotaxus fortunei Hook. var. alpina
Calocedrus macrolepis Kurz var. formosana (Florin) H. L. Li 262, 263
W. C. Cheng & L. K. Fu 248 Cephalotaxus fortunei Hook. var. fortunei 262,
Calocedrus rupestris Aver., Hiep & L. K. Phan 263, 331
246, 250, 251, 330, 1053 Cephalotaxus fortunei Hook. var. globosa
Caryopitys Small 608 S. Y. Hu 263
Caryopitys edulis (Engelm.) Small 608 Cephalotaxus fortunei Hook. var. lanceolata
Caryotaxus Henkel & W. Hochst. 998 (K. M. Feng) Silba 267
Caryotaxus nucifera (L.) Henkel & W. Hochst. 998 Cephalotaxus griffithii Hook. f. 269, 271
Cathaya Chun & Kuang 17, 38, 49, 50, 252 Cephalotaxus hainanensis H. L. Li 262, 264, 270
Cathaya argyrophylla Chun & Kuang 252, 253, Cephalotaxus harringtonii (Knight ex J. Forbes)
331 K. Koch 261, 262, 265, 266, 267, 269, 273, 332
Cedrus Trew 38, 49, 50, 109, 254, 256, 258, 301, 924 Cephalotaxus harringtonii (Knight ex J. Forbes)
Cedrus atlantica (Endl.) Manetti ex Carrière 109, K. Koch var. harringtonii 266, 331
112, 254, 255, 256, 259, 301, 331, 466, 734, 986 Cephalotaxus harringtonii (Knight ex J. Forbes)
Cedrus brevifolia (Hook. f.) A. Henry 260 K. Koch var. nana (Nakai) Rehd. 265, 266
Cephalotaxus harringtonii (Knight ex J. Forbes) Chamaecyparis thyoides (L.) Britton, Sterns &
K. Koch var. sinensis (Rehd. & E. H. Wilson) Poggenb. 274, 282, 332, 582, 597, 671, 753
Rehd. 272 Chamaecyparis thyoides (L.) Britton, Sterns & Pog­
Cephalotaxus harringtonii (Knight ex J. Forbes) genb. subsp. henryae (H. L. Li) E. Murray 283
K. Koch var. wilsoniana (Hayata) Kitam. Chamaecyparis thyoides (L.) Britton, Sterns &
266, 267 Poggenb. var. henryae (H. L. Li) Little 283
Cephalotaxus koreana Nakai 265, 266 Chamaecyparis thyoides (L.) Britton, Sterns &
Cephalotaxus lanceolata K. M. Feng 261, 267 Poggenb. var. thyoides 283
Cephalotaxus latifolia L. K. Fu & R. R. Mill 262, Chrysolarix H. E. Moore 924
268, 269 Chrysolarix amabilis (J. Nelson) H. E. Moore 924
Cephalotaxus mannii Hook. f. 173, 176, 251, 262, Columbea Salisb. 185 1081
264, 269, 270, 271, 332, 984 Columbea angustifolia Bertol. 188
Cephalotaxus nana Nakai 266 Cryptomeria D. Don 34, 46, 47, 284, 285, 392
Cephalotaxus oliveri Mast. 261, 269, 271 Cryptomeria fortunei Hooibr. 284
Cephalotaxus sinensis (Rehd. & E. H. Wilson) Cryptomeria japonica (Thunb. ex L. f.) D. Don 18,
H. L. Li 261, 267, 269, 272 26, 83, 172, 267, 280, 284, 289, 332, 476, 930,
Cephalotaxus sinensis (Rehd. & E. H. Wilson) H. L. 947, 996, 1019, 1055, 1062
Li var. latifolia W. C. Cheng & L. K. Fu 268 Cryptomeria japonica (Thunb. ex L. f.) D. Don var.
Cephalotaxus sinensis (Rehd. & E. H. Wilson) H. L. fortunei (Hooibr.) Henry 284
Li var. wilsoniana (Hayata) L. K. Fu & Nan Cryptomeria japonica (Thunb. ex L. f.) D. Don
Li 267 subsp. sinensis (Miq.) P. D. Sell 284
Cephalotaxus sumatrana Miq. 983 Cryptomeria japonica (Thunb. ex L. f.) D. Don var.
Cephalotaxus wilsoniana Hayata 267 sinensis Miq. 284
Chamaecyparis Spach 35, 46, 47, 266, 274, 291, Cunninghamia R. Br. 34, 46, 249, 286, 287
303, 372, 1028 Cunninghamia konishii Hayata 275, 280, 286, 287,
Chamaecyparis formosensis Matsum. 274, 275, 288, 333, 955, 1059
280, 287, 332, 955 Cunninghamia lanceolata (Lamb.) Hook. 286,
Chamaecyparis funebris (Endl.) Franco 302 287, 288, 289, 333, 374, 473, 475, 707, 925, 1059
Chamaecyparis henryae H. L. Li 283 Cunninghamia sinensis R. Br. 286
Chamaecyparis hodginsii (Dunn) Rushforth 372 Cunninghamia unicanaliculata D. Y. Wang & H. L.
Chamaecyparis lawsoniana (A. Murray bis) Liu 288
Parl. 247, 274, 276, 277, 332, 600, 715, 993 Cunninghamia unicanaliculata D. Y. Wang & H. L.
Chamaecyparis nootkatensis (D. Don) Spach 61, Liu var. pyramidalis D. Y. Wang & H. L. Liu  288
1029, 1030 Cuprespinnata J. Nelson 956
Chamaecyparis obtusa (Siebold & Zucc.) Cuprespinnata disticha (L.) J. Nelson 956
Endl. 83, 274, 275, 277, 278, 281, 285, 930, 947, Cupressaceae Gray 15, 16, 18, 19, 22, 24, 25, 26, 33,
994, 996, 1005, 1019 43, 45, 46, 132, 174, 216, 220, 221, 224, 228, 232,
Chamaecyparis obtusa (Siebold & Zucc.) Endl. 237, 246, 251, 274, 284, 286, 290, 315, 316, 361,
subsp. formosana (Hayata) H. L. Li 280 370, 372, 375, 381, 391, 423, 437, 448, 457, 503,
Chamaecyparis obtusa (Siebold & Zucc.) Endl. var. 508, 512, 514, 525, 536, 543, 606, 793, 852, 922,
formosana (Hayata) Hayata 95, 275, 278, 280, 948, 950, 954, 956, 986, 989, 993, 1020, 1025,
287, 587, 955 1028, 1034, 1035, 1038, 1040, 1041, 1053, 1054,
Chamaecyparis obtusa (Siebold & Zucc.) Endl. var. 1055, 1056, 1057, 1059, 1060, 1062, 1063
obtusa 278, 279, 280 Cupressus L. 19, 24, 35, 45, 46, 48, 290, 291, 301,
Chamaecyparis obtusa (Siebold & Zucc.) Endl. var. 303, 313, 372, 1028
obtusa f. formosana Hayata 280 Cupressus abramsiana C. B. Wolf 305
Chamaecyparis pisifera (Siebold & Zucc.) Endl. Cupressus arizonica Greene 78, 125, 126, 291, 292,
274, 275, 276, 280, 947, 994, 996 333, 566
Chamaecyparis sphaeroidea (Spreng.) Spach 274 Cupressus arizonica Greene var. arizonica 293
Chamaecyparis taiwanensis Masam. & S. Suzuki   280 Cupressus arizonica Greene var. glabra (Sudw.)
­Little 293, 1030
 Cupressus arizonica Greene var. montana (Wig- Cupressus hodginsii Dunn 372
gins) Little 293 Cupressus horizontalis Mill. 301, 313
Cupressus arizonica Greene var. nevadensis Cupressus japonica Thunb. ex L. f. 284
(Abrams) Little 294 Cupressus jiangensis N. Zhao 299
Cupressus arizonica Greene var. revealiana Silba 294 Cupressus juniperoides L. 1021
Cupressus arizonica Greene var. stephensonii Cupressus karnaliensis Silba 315
(C. B. Wolf) Little 294 Cupressus karnaliensis Silba var. mustangensis
Cupressus atlantica Gaussen 302 Silba 315
Cupressus austrotibetica Silba 315 Cupressus lawsoniana A. Murray bis 276
Cupressus bakeri Jeps. 291, 295 Cupressus lindleyi Klotzsch ex Endl. 308, 566
1082 Cupressus bakeri Jeps. subsp. matthewsii C. B. Cupressus lusitanica Mill. 78, 89, 91, 92, 291, 307,
Wolf 295 333, 566, 666, 668, 680, 683, 701, 713, 771, 981,
Cupressus benthamii Endl. 308 1025, 1030
Cupressus cashmeriana Royle ex Carrière 291, Cupressus lusitanica Mill. var. benthamii (Endl.)
296, 297 Carrière 308
Cupressus chengiana S. Y. Hu 291, 297 Cupressus lusitanica Mill. var. hondurensis
Cupressus chengiana S. Y. Hu var. chengiana 298 Silba 308
Cupressus chengiana S. Y. Hu var. jiangensis Cupressus lusitanica Mill. var. lusitanica 308
(N. Zhao) Silba 299 Cupressus lusitanica Mill. subsp. torulosa (D. Don)
Cupressus columnaris J. R. Forst. 196 Silba 315
Cupressus disticha L. 956, 958 Cupressus macnabiana A. Murray bis 291, 309,
Cupressus disticha L. var. imbricaria Nutt. 958 310
Cupressus disticha L. var. nutans Aiton 958 Cupressus macrocarpa Hartw. ex Gordon 290,
Cupressus duclouxiana Hickel 291, 299 291, 311, 333, 749, 1030
Cupressus dupreziana A. Camus 290, 291, 300, Cupressus montana Wiggins 293
301, 333, 1053 Cupressus nevadensis Abrams 294
Cupressus dupreziana A. Camus var. atlantica Cupressus nootkatensis D. Don 1028, 1029
(Gaussen) Silba 301, 302 Cupressus pakistanensis Silba 315
Cupressus dupreziana A. Camus var. dup­ Cupressus pendula Griff. 296, 297
reziana 301 Cupressus pendula Thunb. 296
Cupressus fallax Franco 298 Cupressus pigmaea (Lemmon) Sarg. 304
Cupressus forbesii Jeps. 306 Cupressus sargentii Jeps. 291, 312
Cupressus funebris Endl. 289, 290, 291, 302, 473 Cupressus sempervirens L. 290, 291, 301, 302, 313,
Cupressus gigantea W. C. Cheng & L. K. Fu 316 641, 1057
Cupressus glabra Sudw. 293 Cupressus sempervirens L. var. atlantica (Gaussen)
Cupressus goveniana Gordon 291, 303 Silba 302
Cupressus goveniana Gordon var. abramsiana Cupressus sempervirens L. var. dupreziana (A. Ca-
(C. B. Wolf) Little 305 mus) Silba 301
Cupressus goveniana Gordon var. goveniana 304 Cupressus sempervirens L. subsp. horizontalis (Mill.)
Cupressus goveniana Gordon subsp. pigmaea (Lem- A. Camus 313
mon) A. Camus 304 Cupressus sempervirens L. var. horizontalis (Mill.)
Cupressus goveniana Gordon var. pigmaea Lem- Loudon 313
mon 304 Cupressus sempervirens L. var. pyramidalis auct.  313
Cupressus guadalupensis S. Watson 291, 305, 306 Cupressus stephensonii C. B. Wolf 294
Cupressus guadalupensis S. Watson subsp. forbesii Cupressus thyoides L. 274, 283
(Jeps.) Beauch. 306 Cupressus tongmaiensis Silba 315
Cupressus guadalupensis S. Watson var. forbesii Cupressus tongmaiensis Silba var. ludlowii Silba 315
(Jeps.) Little 306, 333 Cupressus tonkinensis Silba 314, 315
Cupressus guadalupensis S. Watson var. guadalu­ Cupressus tortulosa Griff. 296
pensis 306 Cupressus torulosa D. Don 258, 291, 314, 315, 333,
Cupressus himalaica Silba 296, 297 422
Cupressus torulosa D. Don var. gigantea (W. C. Dacrydium beccarii Parl. var. kinabaluense Cor-
Cheng & L. K. Fu) Farjon 315, 316, 468 ner 348
Cupressus torulosa D. Don var. torulosa 315 Dacrydium beccarii Parl. var. rudens de Laub. 353
Cupressus vietnamensis (Farjon) Silba 1030 Dacrydium beccarii Parl. var. subelatum Corner 354
×Cuprocyparis leylandii (A. B. Jackson & Dallimore) Dacrydium bidwillii Hook. f. ex Kirk 377, 378
Farjon 1030 Dacrydium biforme (Hook.) Pilg. 378
Dacrydium colensoi Hook. 510
Dacrydium comosum Corner 335, 339, 343, 344
D
Dacrydium cornwallianum de Laub. 339, 344
Dacrycarpus (J. J. Bennett) de Laub. 26, 40, 51, 52, Dacrydium cupressinum Sol. ex G. Forst. 147, 322,
317, 319, 320, 343, 347, 351, 365, 367, 524, 547, 812, 335, 337, 339, 345, 346, 380, 498, 505, 507, 549, 1083
884, 953 827, 908, 916, 923
Dacrycarpus cinctus (Pilg.) de Laub. 317, 318, 319, Dacrydium elatum (Roxb.) Wall. ex Hook. 176,
334 251, 336, 338, 339, 346, 347, 659, 695
Dacrycarpus compactus (Wassch.) de Laub. 317, Dacrydium ericoides de Laub. 338, 347, 358
319, 320, 815 Dacrydium falciforme (Parl.) Pilg. 366
Dacrycarpus cumingii (Parl.) de Laub. 317, 320, Dacrydium fitzgeraldii F. Muell. 541
321 Dacrydium fonkii (Phil.) Benth. & Hook. f. 498
Dacrycarpus dacrydioides (A. Rich.) de Dacrydium franklinii Hook. f. 478
Laub. 147, 317, 318, 321, 334, 346, 380, 507, 549, Dacrydium gibbsiae Stapf 326, 336, 338, 348, 349,
827, 908, 916, 923 818, 837
Dacrycarpus expansus de Laub. 317, 322, 323 Dacrydium gracile de Laub. 326, 336, 338, 349,
Dacrycarpus imbricatus (Blume) de Laub. 176, 350, 851
251, 270, 318, 319, 321, 323, 324, 325, 326, 327, Dacrydium guillauminii J. T. Buchholz 336, 338,
334, 349, 350, 355, 365, 868, 943, 984 359, 525, 350, 940, 1058
Dacrycarpus imbricatus (Blume) de Laub. var. cur­ Dacrydium hookerianum (W. Archer) Eichler 542
vulus (Miq.) de Laub. 325 Dacrydium intermedium Kirk 499
Dacrycarpus imbricatus (Blume) de Laub. var. im­ Dacrydium intermedium Kirk var. gracilis Kirk
bricatus 324, 325 499, 500
Dacrycarpus imbricatus (Blume) de Laub. var. patu­ Dacrydium kirkii F. Muell. ex Parl. 379
lus de Laub. 324, 325 Dacrydium laxifolium Hook. f. 500
Dacrycarpus imbricatus (Blume) de Laub. var. ro­ Dacrydium leptophyllum (Wasscher) de Laub. ex
bustus de Laub. 319, 325 Silba 339, 351
Dacrycarpus kinabaluensis (Wassch.) de Laub. Dacrydium lycopodioides Brongn. & Gris 338,
318, 325, 326, 334, 349, 817 352, 356
Dacrycarpus leptophyllus (Wasscher) Gaussen 351 Dacrydium magnum de Laub. 339, 353
Dacrycarpus steupii de Laub. 325 Dacrydium medium de Laub. 339, 353, 354
Dacrycarpus steupii (Wasscher) de Laub. 318, Dacrydium nausoriense de Laub. 339, 354, 355,
326, 327 356, 943
Dacrycarpus vieillardii (Parl.) de Laub. 318, 327, Dacrydium nidulum de Laub. 339, 355, 356, 357,
328, 525 943
Dacrydium Sol. ex G. Forst. 17, 18, 26, 40, 51, 52, Dacrydium nidulum de Laub. var. araucarioides de
321, 324, 337, 338, 339, 347, 351, 360, 365, 366, Laub. 344
367, 524, 537, 539, 547, 812, 868, 884, 890, 953 Dacrydium novoguineense Gibbs 339, 356, 357,
Dacrydium araucarioides Brongn. & Gris 130, 810
164, 208, 242, 328, 335, 337, 338, 340, 341, 344, Dacrydium pancheri Brongn. & Gris 129
359, 525, 871, 1058 Dacrydium papuanum (de Laub.) Whitmore 366
Dacrydium balansae Brongn. & Gris 242, 338, Dacrydium pectinatum de Laub. 148, 338, 339,
340, 341 357, 358
Dacrydium beccarii Parl. 335, 338, 339, 341, 342, Dacrydium pectinatum de Laub. var. robustum de
343, 347, 360 Laub. 357
 Dacrydium pierrei Hickel 346, 347 E
Dacrydium plumosum D. Don 503, 507
Eutacta Link 185
Dacrydium spathoides de Laub. 339, 348, 358
Eutacta muelleri Carrière 207
Dacrydium ×suprinii Nimsch 338, 359
Eutacta pancheri Carrière 525
Dacrydium taxifolium Banks & Sol. ex D. Don 922
Eutassa Salisb. 185
Dacrydium taxoides Brongn. & Gris 368
Eutassa heterophylla Salisb. 200
Dacrydium tetragonum (Hook.) Parl. 516
Dacrydium ustum Vieill. 539
Dacrydium xanthandrum Pilg. 336, 338, 339, 360 F
Dammara (Rumph.) Lam. 143
Falcatifolium de Laub. 40, 51, 52, 337, 347, 363, 365
1084 Dammara australis D. Don 146
Falcatifolium angustum de Laub. 363, 364
Dammara brownii hort. ex Lem. 157
Falcatifolium falciforme (Parl.) de Laub. 148, 350,
Dammara celebica Koord. 150
363, 364, 365, 366, 449, 524, 851
Dammara hypoleuca C. Moore ex Henkel &
Falcatifolium gruezoi de Laub. 343, 363, 365, 366,
W. Hochst. 163
953
Dammara lanceolata Sébert & Pancher 155
Falcatifolium papuanum de Laub. 363, 366, 368,
Dammara loranthifolia Link 143
810
Dammara macrophylla Lindl. 157
Falcatifolium sleumeri de Laub. & Silba 363, 367,
Dammara moorei Lindl. 161
368
Dammara motleyi Parl. 520
Falcatifolium taxoides (Brongn. & Gris) de Laub.
Dammara obtusa Lindl. 157
18, 40, 202, 205, 352, 363, 368, 369, 449, 504,
Dammara ovata C. Moore ex Vieill. 163
539, 540, 938
Dammara robusta C. Moore ex F. Muell. 166
Falcatifolium usan-apuense de Laub. & Silba
Dammara vitiensis Seem. 157
364, 365
Decussocarpus de Laub. 520, 937
Fitzroya Lindl. 37, 46, 48, 370, 371, 607
Decussocarpus comptonii (J. T. Buchholz) de
Fitzroya cupressoides (Molina) I. M. Johnst. 221,
Laub. 937
370, 371, 449, 499, 607, 872, 1057
Decussocarpus falcatus (Thunb.) de Laub. 138
Foetataxus J. Nelson 998
Decussocarpus fleuryi (Hickel) de Laub. 518
Foetataxus montana J. Nelson 998
Decussocarpus gracilior (Pilg.) de Laub. 139
Fokienia A. Henry & H. H. Thomas 35, 46, 47, 372
Decussocarpus mannii (Hook.) de Laub. 141
Fokienia hodginsii (Dunn) A. Henry & H. H. Tho-
Decussocarpus maximus de Laub. 520
mas 176, 251, 262, 270, 372, 373, 374, 450, 519,
Decussocarpus minus (Carrière) de Laub. 938
522, 535, 659, 661, 695, 858, 955, 975
Decussocarpus motleyi (Parl.) de Laub. 520
Frenela Mirb. 224
Decussocarpus nagi (Thunb.) de Laub. 521, 937
Frenela canescens Parl. 226
Decussocarpus nagi (Thunb.) de Laub. var. formo­
Frenela drummondii Parl. 229
sensis (Dummer) Silba 521
Frenela endlicheri Parl. 230
Decussocarpus piresii Silba 940
Frenela muelleri Parl. 234
Decussocarpus rospigliosii (Pilg.) de Laub. 941
Frenela robusta A. Cunn. ex Endl. var. microcarpa
Decussocarpus vitiensis (Seem.) de Laub. 942
Benth. 227
Decussocarpus wallichianus (C. Presl) de Laub. 523
Frenela roei Endl. 240
Diselma Hook. f. 37, 46, 48, 361
Frenela sulcata Parl. 241
Diselma archeri Hook. f. 19, 217, 219, 336, 361, 362,
Frenela triquetra Spach 224
516, 543, 545, 852
Frenela verrucosa A. Cunn. ex Endl. 243
Dolophyllum Salisb. 996
Dolophyllum dolabrata (L. f.) Salisb. 996
Dombeya Lam. 185 G
Ducampopinus A. Chev. 608
Gigantabies J. Nelson 948
Ducampopinus krempfii (Lecomte) A. Chev. 608,
Gigantabies taxifolia J. Nelson 948
693, 695
Glyptostrobus Endl. 18, 35, 46, 47, 375, 376
Glyptostrobus lineatus (Poir.) Druce 375
Glyptostrobus pensilis (Staunton ex D. Don) Juniperus ashei J. T. Buchholz var. ashei 387
K. Koch 375, 376, 450 Juniperus ashei J. T. Buchholz var. ovata R. P.
­Adams 387
Juniperus ashei J. T. Buchholz var. saltillensis
H
(M. T. Hall) Silba 457
Halocarpus Quinn 40, 51, 52, 337, 377 Juniperus baimashanensis Y. F. Yu & L. K. Fu 463
Halocarpus bidwillii (Hook. f. ex Kirk) Quinn Juniperus barbadensis L. 384, 388, 389, 459
322, 377, 378, 379, 450, 502, 870 Juniperus barbadensis L. var. barbadensis 388
Halocarpus biformis (Hook.) Quinn 377, 378, 379, Juniperus barbadensis L. var. lucayana (Britton)
450, 502, 505, 550, 827 R. P. Adams 389
Halocarpus kirkii (F. Muell. ex Parl.) Quinn 346, Juniperus barbadensis L. subsp. saxicola (Britton & 1085
377, 379, 380, 507, 510, 827, 923 P. Wilson) Borhidi 459
Hesperocyparis Bartel & R. A. Price 290, 291 Juniperus barbadensis L. var. saxicola (Britton &
Hesperocyparis abramsiana (C. B. Wolf) Bartel 305 P. Wilson) Silba 459
Hesperocyparis arizonica (Greene) Bartel 293 Juniperus barbadensis L. subsp. urbaniana (Pilg. &
Hesperocyparis bakeri (Jeps.) Bartel 295 Ekman) Borhidi 420
Hesperocyparis benthamii (Endl.) Bartel 308 Juniperus barbadensis L. var. urbaniana (Pilg. &
Hesperocyparis forbesii (Jeps.) Bartel 306 ­Ekman) Silba 420
Hesperocyparis glabra (Sudw.) Bartel 293 Juniperus bermudiana L. 384, 389, 390
Hesperocyparis goveniana (Gordon) Bartel 304 Juniperus blancoi Martínez 384, 390, 391, 461
Hesperocyparis guadalupensis (S. Watson) Bar- Juniperus blancoi Martínez var. blancoi 391
tel 306 Juniperus blancoi Martínez var. huehuentensis
Hesperocyparis lusitanica (Mill.) Bartel 308 R. P. Adams 391
Hesperocyparis macnabiana (A. Murray bis) Bar- Juniperus blancoi Martínez var. mucronata
tel 309 (R. P. Adams) Farjon 391
Hesperocyparis macrocarpa (Hartw. ex Gordon) Juniperus brevifolia (Seub.) Antoine 382, 392, 394
Bartel 290, 311 Juniperus californica Carrière 292, 393, 450, 712,
Hesperocyparis montana (Wiggins) Bartel 293 748, 749
Hesperocyparis nevadensis (Abrams) Bartel 294 Juniperus californica Carrière f. lutheyana J. T.
Hesperocyparis pigmaea (Lemmon) Bartel 304 ­Howell & Twisselm. 393
Hesperocyparis sargentii (Jeps.) Bartel 312 Juniperus californica Carrière subsp. osteosperma
Hesperocyparis stephensonii (C. B. Wolf) Bartel  294 (Torr.) E. Murray 429
Hesperopeuce (Engelm.) Lemmon 1007 Juniperus canariensis Guyot & Mathou 434
Hesperopeuce mertensiana (Bong.) Lemmon 1007 Juniperus carinata (Y. F. Yu & L. K. Fu) R. P.
Hesperopeuce mertensiana (Bong.) Rydb. 1018 ­Adams 437
×Hesperotsuga C. N. Page 1007 Juniperus cedrus Webb & Berthel. 382, 394
×Hesperotsuga jeffreyi (A. Henry) C. N. Page 1007, Juniperus cedrus Webb & Berthel. subsp. maderensis
1018 (Menezes) Rivas Mart. et al. 394
Juniperus chengii L. K. Fu & Y. F. Yu 437
Juniperus chinensis L. 384, 395, 396, 447, 450, 459,
J
461, 469, 470, 1059
Juniperus L. 19, 23, 24, 26, 35, 46, 47, 89, 96, 134, Juniperus chinensis L. var. arenaria E. H. Wilson  448
290, 308, 381, 393, 398, 412, 416, 418, 435, 458, Juniperus chinensis L. var. chinensis 396
463, 467, 484, 487, 514, 670, 673, 700, 701, 765, Juniperus chinensis L. var. procumbens Siebold ex
1028 Endl. 439
Juniperus africana (Maire) Villar 466 Juniperus chinensis L. var. sargentii A. Henry 396
Juniperus angosturana R. P. Adams 383, 385 Juniperus chinensis L. var. tsukusiensis (Masam.)
Juniperus arenaria (E. H. Wilson) Florin 448 Masam. 397
Juniperus arizonica (R. P. Adams) R. P. Adams Juniperus chinensis Roxb. 859
384, 385 Juniperus coahuilensis (Martínez) Gaussen ex
Juniperus ashei J. T. Buchholz 383, 386, 469, 751 R. P. Adams 384, 386, 397, 398
 Juniperus coahuilensis (Martínez) Gaussen ex Juniperus deppeana Steud. var. zacatecensis Mar-
R. P. Adams var. arizonica R. P. Adams 385 tínez 407
Juniperus comitana Martínez 383, 398, 399 Juniperus depressa (Pursh) Raf. 401
Juniperus communis L. 27, 103, 109, 255, 381, 382, Juniperus distans Florin 467, 468
399, 400, 403, 418, 421, 451 Juniperus drupacea Labill. 314, 381, 382, 407, 408,
Juniperus communis L. var. charlottensis R. P. Adams 409, 416, 1054
402, 403 Juniperus durangensis Martínez 384, 410
Juniperus communis L. var. communis 400, 451 Juniperus ekmanii Florin 420
Juniperus communis L. subsp. depressa (Pursh) Juniperus elata Roxb. 346
Franco 401 Juniperus erectopatens (W. C. Cheng & L. K. Fu)
1086 Juniperus communis L. var. depressa Pursh 401 R. P. Adams 396
Juniperus communis L. var. megistocarpa Fernald Juniperus erythrocarpa Cory 435
& H. St. John 402 Juniperus erythrocarpa Cory var. coahuilensis Mar-
Juniperus communis L. var. montana Aiton 400, tínez 397
402 Juniperus excelsa M.-Bieb. 72, 259, 314, 384, 409,
Juniperus communis L. var. nana (Willd.) 411, 412, 416, 431, 641
Baumg. 400 Juniperus excelsa M.-Bieb. subsp. excelsa 412
Juniperus communis L. var. nipponica (Maxim.) Juniperus excelsa M.-Bieb. subsp. polycarpos
E. H. Wilson 100, 402, 403 (K. Koch) Takht. 411, 412, 413, 462
Juniperus communis L. subsp. pygmaea (K. Koch) Juniperus excelsa M.-Bieb. var. polycarpos (K. Koch)
Imkhan. 402 Silba 412
Juniperus communis L. var. saxatilis Pall. 400, Juniperus excelsa M.-Bieb. subsp. polycarpos
402, 451, 462, 484 (K. Koch) Takht. var. pendula (Mulk.) Im-
Juniperus compacta (Martínez) R. P. Adams 427 khan. 412
Juniperus conferta Parl. 446 Juniperus excelsa M.-Bieb. subsp. seravschanica
Juniperus convallium Rehd. & E. H. Wilson 383, (Kom.) Imkhan. 412
403, 404, 459, 467 Juniperus excelsa M.-Bieb. subsp. turcomanica
Juniperus convallium Rehd. & E. H. Wilson var. (B. Fedtsch.) Imkhan. 412
convallium 404 Juniperus flaccida Schltdl. 384, 385, 405, 414, 415,
Juniperus convallium Rehd. & E. H. Wilson var. mi­ 451, 623, 680, 737, 744, 1053
crosperma (W. C. Cheng & L. K. Fu) Silba 404 Juniperus flaccida Schltdl. var. flaccida 415
Juniperus coxii A. B. Jacks. 444 Juniperus flaccida Schltdl. var. martinezii (Pérez de
Juniperus davurica Pall. 457 la Rosa) Silba 415
Juniperus davurica Pall. subsp. maritima Urus­ Juniperus flaccida Schltdl. var. poblana Mar­
sov 457 tínez 415
Juniperus deltoides R. P. Adams 432 Juniperus foetidissima Willd. 259, 314, 384, 409,
Juniperus deppeana Steud. 78, 384, 405, 410, 415, 412, 416, 431
418, 451, 623, 668, 1053 Juniperus formosana Hayata 75, 289, 382, 417
Juniperus deppeana Steud. var. deppeana 406, Juniperus formosana Hayata var. mairei (Lemée &
418 Lév.) R. P. Adams & C. F. Hsieh 417
Juniperus deppeana Steud. var. gamboana (Mar- Juniperus gamboana Martínez 418
tínez) R. P. Adams 418 Juniperus gaussenii W. C. Cheng 470, 471
Juniperus deppeana Steud. var. pachyphlaea (Torr.) Juniperus gracilior Pilg. 384, 419
Martínez 292, 406 Juniperus gracilior Pilg. var. ekmanii (Florin)
Juniperus deppeana Steud. var. patoniana (Mar- R. P. Adams 420
tínez) Zanoni 406 Juniperus gracilior Pilg. var. gracilior 419
Juniperus deppeana Steud. var. robusta Martínez Juniperus gracilior Pilg. var. urbaniana (Pilg. &
391, 406, 418 ­Ekman) R. P. Adams 420
Juniperus deppeana Steud. var. sperryi Correll  407 Juniperus grandis R. P. Adams 429
Juniperus deppeana Steud. f. zacatensis (Martínez) Juniperus horizontalis Moench 384, 420, 421, 422,
R. P. Adams 407 469
Juniperus horizontalis Moench subsp. hamptonensis Juniperus oxycedrus L. var. fastigiata Jovan. 432
Silba 420 Juniperus oxycedrus L. subsp. macrocarpa (Sibth.
Juniperus horizontalis Moench subsp. neopangaea & Sm.) Ball 409, 431, 432, 446, 451
Silba 420 Juniperus oxycedrus L. subsp. maderensis Men-
Juniperus indica Bertol. 19, 383, 422, 463 ezes 394
Juniperus indica Bertol. var. caespitosa Farjon Juniperus oxycedrus L. subsp. oxycedrus 431
422, 423 Juniperus oxycedrus L. var. oxycedrus f. parvifolia
Juniperus indica Bertol. var. indica 422, 423 Novák 431
Juniperus jaliscana Martínez 384, 423 Juniperus oxycedrus L. subsp. oxycedrus var.
Juniperus komarovii Florin 383, 424 spilinanus Yaltirik, Eliçin & Terzioglu 432
Juniperus lucayana Britton 389 Juniperus oxycedrus L. var. parvifolia (Novák) 1087
Juniperus lutchuensis Koidz. 465 ­Jovan. 431
Juniperus macrocarpa Sibth. & Sm. 432 Juniperus oxycedrus L. subsp. transtagana Franco
Juniperus macropoda Boiss. 412, 462 431, 433
Juniperus mairei Lemée & Lév 417 Juniperus oxycedrus L. var. transtagana (Franco)
Juniperus maritima R. P. Adams 460, 461 Silba 433
Juniperus martinezii Pérez de la Rosa 415 Juniperus patoniana Martínez 406
Juniperus media V. D. Dmitriev 396, 461 Juniperus ×pfitzeriana (Späth) Schmidt 396
Juniperus microsperma (W. C. Cheng & L. K. Fu) Juniperus phoenicea L. 301, 314, 381, 383, 433, 986
R. P. Adams 404 Juniperus phoenicea L. subsp. eu-mediterranea
Juniperus monosperma (Engelm.) Sarg. 383, 386, P. Lebreton & S. Thivend 434
425, 426, 670, 751 Juniperus phoenicea L. subsp. phoenicea 434, 452
Juniperus monosperma (Engelm.) Sarg. var. gracilis Juniperus phoenicea L. subsp. turbinata (Guss.)
Martínez 385 Nyman 434
Juniperus monosperma (Engelm.) Sarg. var. pin­ Juniperus pinchotii Sudw. 383, 386, 387, 435
chotii (Sudw.) Melle 435 Juniperus pinchotii Sudw. var. erythrocarpa (Cory)
Juniperus montana (Aiton) Lindl. & Gordon 402 Silba 435
Juniperus monticola Martínez 384, 427 Juniperus pingii W. C. Cheng 383, 436, 459
Juniperus monticola Martínez var. monticola f. com­ Juniperus pingii W. C. Cheng var. carinata Y. F. Yu &
pacta Martínez 427 L. K. Fu 437
Juniperus morrisonicola Hayata 462 Juniperus pingii W. C. Cheng var. chengii (L. K. Fu
Juniperus mucronata R. P. Adams 391 & Y. F. Yu) Farjon 437
Juniperus navicularis Gand. 433 Juniperus pingii W. C. Cheng var. miehei Farjon  437
Juniperus nipponica Maxim. 402 Juniperus pingii W. C. Cheng var. pingii 436
Juniperus occidentalis Hook. 99, 383, 386, 428, Juniperus pingii W. C. Cheng var. wilsonii (Rehd.)
430, 451, 638, 690, 712, 757, 1017 Silba 436, 437, 438, 444, 463, 464
Juniperus occidentalis Hook. subsp. australis Juniperus poblana (Martínez) R. P. Adams 415
Vasek 429 Juniperus polycarpos K. Koch 412, 462
Juniperus occidentalis Hook. var. australis (Vasek) Juniperus polycarpos K. Koch var. pendula
P. Lebreton & N. H. Holmgren 428, 429 Mulk. 412
Juniperus occidentalis Hook. var. monosperma Juniperus polycarpos K. Koch var. seravschanica
Engelm. 425 (Kom.) Kitam. 412
Juniperus occidentalis Hook. var. occidentalis Juniperus polycarpos K. Koch var. turcomanica
428, 429 (B. Fedtsch.) R. P. Adams 412
Juniperus osteosperma (Torr.) Little 383, 386, 425, Juniperus procera Hochst. ex Endl. 140, 384, 438,
429, 430, 461, 670, 712, 1061 439, 865
Juniperus oxycedrus L. 382, 394, 409, 410, 412, Juniperus procumbens (Siebold ex Endl.) Miq.
416, 430 383, 439
Juniperus oxycedrus L. subsp. badia (H. Gay) Juniperus przewalskii Kom. 383, 440, 441
­Debeaux 431, 432 Juniperus pseudosabina Fisch. & C. A. Mey. 383,
Juniperus oxycedrus L. var. badia H. Gay 432 411, 422, 441, 448, 452, 462, 598
 Juniperus pseudosabina Fisch. & C. A. Mey. var. Juniperus squamata Buch.-Ham. ex D. Don 77,
turkestanica (Kom.) Silba 441 121, 258, 383, 414, 417, 422, 436, 438, 459, 462,
Juniperus pygmaea K. Koch 402 463, 464, 470, 495
Juniperus recurva Buch.-Ham. ex D. Don 382, Juniperus squamata Buch.-Ham. ex D. Don var.
436, 438, 442, 443, 444, 463 fargesii Rehd. & E. H. Wilson 463
Juniperus recurva Buch.-Ham. ex D. Don var. coxii Juniperus squamata Buch.-Ham. ex D. Don var.
(A. B. Jacks.) Melville 444, 445 hongxiensis Y. F. Yu & L. K. Fu 463
Juniperus recurva Buch.-Ham. ex D. Don var. re­ Juniperus squamata Buch.-Ham. ex D. Don var.
curva 444 morrisonicola (Hayata) H. L. Li & H. Keng
Juniperus rigida Siebold & Zucc. 382, 440, 445, 95, 462, 587
1088 446, 584, 794 Juniperus squamata Buch.-Ham. ex D. Don var.
Juniperus rigida Siebold & Zucc. subsp. conferta parvifolia Y. F. Yu & L. K. Fu 463
(Parl.) Kitam. 445, 446 Juniperus squamata Buch.-Ham. ex D. Don f. wilso­
Juniperus rigida Siebold & Zucc. var. conferta (Parl.) nii Rehd. 437
Patschke 446 Juniperus standleyi Steyerm. 385, 399, 464, 683
Juniperus rigida Siebold & Zucc. subsp. litoralis Juniperus taxifolia Hook. & Arn. 382, 465
Urussov 446 Juniperus tetragona Schltdl. var. osteosperma
Juniperus rigida Siebold & Zucc. var. litoralis Torr. 429
(Urussov) Kozhevnikova 446 Juniperus texensis Melle 435
Juniperus rigida Siebold & Zucc. subsp. nipponica Juniperus thurifera L. 385, 466
(Maxim.) Franco 402 Juniperus thurifera L. subsp. africana (Maire) Romo
Juniperus rigida Siebold & Zucc. subsp. rigida  446 & Borantinsky 466
Juniperus sabina L. 381, 384, 396, 411, 442, 447, Juniperus thurifera L. var. africana Maire 466
457, 459, 461, 462 Juniperus tianschanica Sumnev. 461
Juniperus sabina L. var. arenaria (E. H. Wilson) Juniperus tibetica Kom. 383, 404, 467
Farjon 448 Juniperus tsukusiensis Masam. 397
Juniperus sabina L. var. davurica (Pall.) Farjon Juniperus turbinata Guss. 434
105, 448, 457, 515 Juniperus turbinata Guss. subsp. canariensis (Guyot
Juniperus sabina L. var. erectopatens (W. C. Cheng & & Mathou) Rivas-Martínez et al. 434
L. K. Fu) Y. F. Yu & L. K. Fu 396 Juniperus turcomanica B. Fedtsch. 412
Juniperus sabina L. var. mongolensis R. P. Adams 448 Juniperus turkestanica Kom. 441
Juniperus sabina L. var. monosperma C. Y. Yang 448 Juniperus urbaniana Pilg. & Ekman 420
Juniperus sabina L. var. sabina 448, 452 Juniperus uvifera D. Don 606
Juniperus sabina L. var. yulinensis (T. C. Chang & Juniperus virginiana L. 384, 390, 421, 468, 469,
C. G. Chen) Y. F. Yu & L. K. Fu 448 1053
Juniperus saltillensis M. T. Hall 384, 415, 457, 458 Juniperus virginiana L. subsp. silicicola (Small)
Juniperus saltuaria Rehd. & E. H. Wilson 383, E. Murray 470
404, 458, 459, 463 Juniperus virginiana L. var. silicicola (Small)
Juniperus sargentii (A. Henry) Takeda ex Nakai 396 E. Murray 470
Juniperus saxicola Britton & P. Wilson 384, 459 Juniperus virginiana L. var. virginiana 453, 469,
Juniperus scopulorum Sarg. 384, 425, 430, 460, 470
461, 469 Juniperus wallichiana Hook. f. & Thomson ex
Juniperus semiglobosa Regel 384, 396, 411, 442, E. Brandis 121, 422, 423, 602
452, 453, 461, 462, 463
Juniperus seravschanica Kom. 412
K
Juniperus sibirica Burgsd. 402, 403
Juniperus silicicola (Small) L. H. Bailey 469, 470, Keteleeria Carrière 38, 49, 50, 249, 472, 473, 533,
1053 924, 1055
Juniperus sphaerica Lindl. 396 Keteleeria calcarea W. C. Cheng & L. K. Fu 474
Keteleeria davidiana (Bertrand) Beissn. 251, 453,
472, 473, 707
Keteleeria davidiana (Bertrand) Beissn. var. calca­ Larix decidua Mill. var. decidua 454, 483
rea (W. C. Cheng & L. K. Fu) Silba 474 Larix decidua Mill. var. polonica (Racib. ex Wóy-
Keteleeria davidiana (Bertrand) Beissn. var. dav­ cicki) Ostenf. & Syrach 483
idiana 474 Larix decidua Mill. subsp. sibirica (Ledeb.)
Keteleeria davidiana (Bertrand) Beissn. subsp. for­ Domin 496
mosana (Hayata) E. Murray 474 Larix decidua Mill. var. sibirica (Ledeb.) Regel   496
Keteleeria davidiana (Bertrand) Beissn. var. for­ Larix ×eurolepis Henry 489
mosana (Hayata) Hayata 474 Larix gmelinii (Rupr.) Kuzen. 17, 105, 120, 457,
Keteleeria davidiana (Bertrand) Beissn. var. pubes­ 480, 481, 483, 484, 586, 590, 746, 759, 978
cens (W. C. Cheng & L. K. Fu) Silba 474 Larix gmelinii (Rupr.) Kuzen. f. genhensis (S. Y. Li &
Keteleeria evelyniana Mast. 251, 472, 474, 475, Adair) L. K. Fu & Nan Li 485 1089
783, 984 Larix gmelinii (Rupr.) Kuzen. var. genhensis S. Y. Li
Keteleeria evelyniana Mast. var. hainanensis (Chun & Adair 485
& Tsiang) Silba 474 Larix gmelinii (Rupr.) Kuzen. var. gmelinii 485
Keteleeria evelyniana Mast. var. pendula J. R. Xue Larix gmelinii (Rupr.) Kuzen. subsp. japonica
474 (­Maxim. ex Regel) E. Murray 485
Keteleeria fabri Mast. 79 Larix gmelinii (Rupr.) Kuzen. var. japonica
Keteleeria formosana Hayata 473, 474 (­Maxim. ex Regel) Pilg. 117, 485
Keteleeria fortunei (A. Murray bis) Carrière 262, Larix gmelinii (Rupr.) Kuzen. var. olgensis (A. Hen-
472, 475, 476, 477, 522 ry) Ostenf. & Syrach 93, 485, 576
Keteleeria fortunei (A. Murray bis) Carrière var. Larix gmelinii (Rupr.) Kuzen. f. pendula (D. S. Zhang
oblonga (W. C. Cheng & L. K. Fu) L. K. Fu & & Y. M. Chen) L. K. Fu & Nan Li 486
Nan Li 475 Larix gmelinii (Rupr.) Kuzen. subsp. principis-­
Keteleeria fortunei (A. Murray bis) Carrière var. rupprechtii (Mayr) E. Murray 486
xerophila (J. R. Xue & S. H. Hao) Silba 474 Larix gmelinii (Rupr.) Kuzen. var. principis-­
Keteleeria hainanensis Chun & Tsiang 472, 474 rupprechtii (Mayr) Pilg. 454, 481, 486
Keteleeria oblonga W. C. Cheng & L. K. Fu 475 Larix griffithiana hort. ex Carrière 77, 486, 487,
Keteleeria pubescens W. C. Cheng & L. K. Fu 472, 602
474 Larix griffithii Hook. f. 445, 480, 481, 486, 487,
Keteleeria xerophila J. R. Xue & S. H. Hao 474 1014
Larix griffithii Hook. f. var. griffithii 454, 487, 488
Larix griffithii Hook. f. var. mastersiana (Rehd. &
L
E. H. Wilson) Silba 491
Lagarostrobos Quinn 40, 51, 52, 337, 478, 510 Larix griffithii Hook. f. var. speciosa (W. C. Cheng
Lagarostrobos colensoi (Hook.) Quinn 510 & Y. W. Law) Farjon 488
Lagarostrobos franklinii (Hook. f.) Quinn 453, Larix himalaica W. C. Cheng & L. K. Fu 495
478, 510 Larix kaempferi (Lamb.) Carrière 94, 123, 454,
Laricopsis Kent 924 480, 481, 488, 489, 557, 577, 604, 978, 994, 1013
Laricopsis kaempferi (Lindl.) Kent 924 Larix kongboensis R. R. Mill 487
Larix Mill. 15, 17, 38, 49, 50, 263, 403, 423, 436, Larix laricina (Du Roi) K. Koch 480, 481, 489,
448, 463, 480, 569, 574, 581, 633, 765, 924 582, 639, 991, 1009
Larix alaskensis W. F. Wight 489 Larix laricina (Du Roi) K. Koch var. alaskensis
Larix amabilis J. Nelson 924 (W. F. Wight) Raup 489
Larix chinensis Beissn. 495 Larix lyallii Parl. 454, 480, 481, 490, 491, 493
Larix ×czekanowskii Szafer 480, 481 Larix ×marschlinsii Coaz 489
Larix dahurica Turcz. ex Trautv. 485 Larix mastersiana Rehd. & E. H. Wilson 480, 481,
Larix dahurica Turcz. ex Trautv. var. japonica 491, 492
­Maxim. ex Regel 485 Larix occidentalis Nutt. 88, 113, 480, 481, 492, 493,
Larix decidua Mill. 25, 454, 480, 481, 482, 486, 568, 715, 993
489, 497, 555, 649 Larix olgensis A. Henry 485
Larix decidua Mill. var. carpatica Domin 483 Larix polonica Racib. ex Wóycicki 483
 Larix potaninii Batalin 70, 79, 82, 84, 114, 122, Manoao colensoi (Hook.) Molloy 322, 346, 378,
445, 459, 480, 481, 491, 493, 494, 495, 496, 562, 455, 507, 510, 511, 549, 827, 916, 923
563, 578, 595, 955, 1015 Margbensonia A. V. Bobrov & Melikyan 795
Larix potaninii Batalin var. chinensis (Beissn.) Margbensonia archboldii (N. E. Gray) A. V. Bobrov
L. K. Fu & Nan Li 495 & Melikyan 810
Larix potaninii Batalin var. himalaica (W. C. Cheng Margbensonia atjehense (Wasscher) A. V. Bobrov &
& L. K. Fu) Farjon & Silba 495 Melikyan 811
Larix potaninii Batalin var. macrocarpa Y. W. Law Margbensonia chingiana (N. E. Gray) A. V. Bobrov
496 & Melikyan 821
Larix potaninii Batalin var. potaninii 495 Margbensonia degeneri (N. E. Gray) A. V. Bobrov &
1090 Larix principis-rupprechtii Mayr 486 Melikyan 869
Larix principis-rupprechtii Mayr var. pendula Margbensonia disperma (C. T. White) A. V. Bobrov
D. S. Zhang & Y. M. Chen 486 & Melikyan 829
Larix russica (Endl.) Sabine ex Trautv. 496 Margbensonia drouyniana (F. Muell.) A. V. Bobrov
Larix sibirica Ledeb. 120, 480, 481, 482, 496, 497, & Melikyan 830
759 Margbensonia elata (R. Br. ex Endl.) A. V. Bobrov &
Larix speciosa W. C. Cheng & Y. W. Law 488 Melikyan 832
Lepidothamnus Phil. 18, 40, 51, 52, 337, 498, 499 Margbensonia koordersii (Pilg. ex Koord. & Vale-
Lepidothamnus fonkii Phil. 370, 455, 498, 499, ton) A. V. Bobrov & Melikyan 890
607 Margbensonia macrophylla (Thunb.) A. V. Bobrov &
Lepidothamnus intermedius (Kirk) Quinn 498, Melikyan 795, 858
499, 500 Margbensonia maki (Siebold & Zucc. ex Endl.)
Lepidothamnus laxifolius (Hook. f.) Quinn 498, A. V. Bobrov & Melikyan 859
500, 501, 502 Margbensonia neriifolia (D. Don) A. V. Bobrov &
Leucopitys Nieuwl. 608 Melikyan 868
Leucopitys strobus (L.) Nieuwl. 608 Margbensonia philippinense (Foxw.) A. V. Bobrov &
Libocedrus Endl. 19, 36, 46, 48, 220, 503, 606 Melikyan 890
Libocedrus arfakensis Gibbs 537 Margbensonia polystachya (R. Br. ex Endl.) A. V. Bo-
Libocedrus austrocaledonica Brongn. & Gris 503, brov & Melikyan 882
504 Margbensonia ridleyi (Wasscher) A. V. Bobrov &
Libocedrus bidwillii Hook. f. 455, 503, 505, 508, Melikyan 886
550, 827 Margbensonia rumphii (Blume) A. V. Bobrov &
Libocedrus chevalieri J. T. Buchholz 503, 506 ­Melikyan 890
Libocedrus chilensis (D. Don) Endl. 220 Margbensonia spinulosa (Sm.) A. V. Bobrov &
Libocedrus decurrens Torr. 246 ­Melikyan 899
Libocedrus decurrens Torr. var. columnaris Beissn. Margbensonia thevetiifolia (Blume) A. V. Bobrov &
247 Melikyan 882
Libocedrus formosana Florin 248 Marywildea A. V. Bobrov & Melikyan 185
Libocedrus macrolepis (Kurz) Benth. & Hook. f.   249 Marywildea bidwillii (Hook.) A. V. Bobrov &
Libocedrus papuana F. Muell. 536, 537 ­Melikyan 185, 194
Libocedrus papuana F. Muell. var. arfakensis (Gibbs) Metadacrydium Baum.-Bod. 337
de Laub. 537 Metadacrydium araucarioides (Brongn. & Gris)
Libocedrus plumosa (D. Don) Sarg. 455, 503, 506, Baum.-Bod. 337, 340
507 Metadacrydium balansae (Brongn. & Gris) Baum.-
Libocedrus uvifera (D. Don) Pilg. 606 Bod. 340
Libocedrus yateensis Guillaumin 508 Metasequoia Hu & W. C. Cheng 15, 18, 34, 46, 512,
513, 922
Metasequoia glyptostroboides Hu & W. C. Cheng
M
15, 455, 512, 1053, 1054
Manoao Molloy 40, 51, 52, 337, 510 Metasequoia glyptostroboides Hu & W. C. Cheng
var. caespitosa Y. H. Long & Y. Wu 512
Microbiota Kom. 36, 46, 47, 514, 793 Nothocallitris neocaledonica (Dummer) A. V. Bo-
Microbiota decussata Kom. 19, 455, 514, 515, 1057 brov & Melikyan 235
Microcachrys Hook. f. 18, 39, 40, 51, 52, 361, 516 Nothocallitris sulcata (Parl.) A. V. Bobrov &
Microcachrys W. Archer 361 ­Melikyan 224, 241
Microcachrys tetragona (Hook.) Hook. f. 18, 361, Nothotaxus Florin 927
362, 456, 516, 543, 852 Nothotaxus chienii (W. C. Cheng) Florin 927
Microstrobos J. Garden & L. A. S. Johnson 541 Nothotsuga Hu ex C. N. Page 39, 49, 50, 533, 924,
Microstrobos fitzgeraldii (F. Muell.) J. Garden & 1007
L. A. S. Johnson 541 Nothotsuga longibracteata (W. C. Cheng) Hu ex
Microstrobos niphophilus J. Garden & L. A. S. John- C. N. Page 253, 374, 533, 534, 535, 625
son 542 1091
Myrica nagi Thunb. 518, 521, 937
O
Octoclinis F. Muell. 224
N
Octoclinis macleayana F. Muell. 224, 231
Nageia Gaertn. 18, 26, 39, 51, 169, 324, 366, 518,
524, 844
P
Nageia amara (Blume) F. Muell. 952
Nageia argotaenia (Hance) Kuntze 169 Pachylepis Brongn. 1020
Nageia comptonii (J. T. Buchholz) de Laub. 937 Pachylepis cupressoides (L.) Brongn. 1020
Nageia falcata (Thunb.) Kuntze 138 Papuacedrus H. L. Li 36, 46, 48, 320, 536, 537
Nageia fleuryi (Hickel) de Laub. 173, 251, 270, 456, Papuacedrus arfakensis (Gibbs) H. L. Li 537
518, 519, 522, 975 Papuacedrus papuana (F. Muell.) H. L. Li 19, 319,
Nageia formosensis (Dummer) C. N. Page 521, 522 320, 323, 327, 536, 625, 810, 826, 1068
Nageia latifolia (Wall.) Gordon 523 Papuacedrus papuana (F. Muell.) H. L. Li var.
Nageia mannii (Hook. f.) Kuntze 141 ­arfakensis (Gibbs) R. J. Johns 536, 537
Nageia maxima (de Laub.) de Laub. 518, 520, 862 Papuacedrus papuana (F. Muell.) H. L. Li var.
Nageia minor Carrière 938 ­papuana 536, 537, 625, 1057
Nageia motleyi (Parl.) de Laub. 518, 520, 521 Parasitaxus de Laub. 41, 51, 52, 337, 539
Nageia nagi (Thunb.) Kuntze 262, 456, 518, 519, Parasitaxus usta (Vieill.) de Laub. 18, 369, 539,
521, 522, 523, 1005 540, 625
Nageia nagi (Thunb.) Kuntze var. formosensis Parolinia Endl. 1020
(Dummer) Silba 521 Pherosphaera W. Archer 41, 51, 52, 541
Nageia nagi (Thunb.) Kuntze var. koshuensis Pherosphaera fitzgeraldii (F. Muell.) Hook. f.
(Kaneh.) D. Z. Fu 522 541, 542
Nageia nankoensis (Hayata) R. R. Mill 521, 522 Pherosphaera hookeriana Hook. f. 516
Nageia pancheri (Brongn. & Gris) Kuntze 129 Pherosphaera hookeriana W. Archer 217, 219, 362,
Nageia piresii (Silba) de Laub. 940 516, 541, 542, 543, 545, 625, 852
Nageia rospigliosii (Pilg.) de Laub. 941 Pherosphaera niphophila (J. Garden & L. A. S. John-
Nageia usta (Vieill.) Kuntze 539 son) Florin 542
Nageia vitiensis (Seem.) Kuntze 942 Phyllocladaceae Bessey 16, 19, 22, 24, 33, 37, 43,
Nageia wallichiana (Presl) Kuntze 15, 22, 148, 175, 48, 49, 544, 1054, 1058, 1059, 1060, 1062
176, 270, 343, 350, 365, 366, 456, 518, 520, 521, Phyllocladus Rich. ex Mirb. 37, 48, 49, 324, 347,
523, 524, 844, 851, 984, 1032 365, 367, 537, 544, 547, 548, 549, 812
Neocallitropsis Florin 37, 46, 48, 525 Phyllocladus alpinus Hook. f. 549, 550
Neocallitropsis araucarioides (R. H. Compton) Phyllocladus aspleniifolius (Labill.) Hook. f. 217,
Florin 525 219, 362, 544, 545, 548, 549, 625
Neocallitropsis pancheri (Carrière) de Laub. 242, Phyllocladus aspleniifolius (Labill.) Hook. f. var.
340, 456, 504, 508, 525 alpinus (Hook. f.) H. Keng 550
Nothocallitris A. V. Bobrov & Melikyan 224 Phyllocladus billardieri Mirb. 544
Phyllocladus glaucus Kirk 547, 548
 Phyllocladus hypophyllus Hook. f. 324, 326, 349, Picea asperata Mast. var. aurantiaca (Mast.)
537, 544, 546, 547, 626, 817, 826, 837, 881, 953 Boom 561
Phyllocladus toatoa Molloy 544, 547, 548, 549 Picea asperata Mast. var. notabilis Rehd. &
Phyllocladus trichomanoides D. Don 147, 322, E. H. Wilson 553, 560
346, 507, 544, 548, 549, 827, 868, 908, 923, 1068 Picea asperata Mast. var. ponderosa Rehd. &
Phyllocladus trichomanoides D. Don var. alpinus E. H. Wilson 553, 561
(Hook. f.) Parl. 378, 379, 505, 550, 626, 827, Picea asperata Mast. var. retroflexa (Mast.) W. C.
870 Cheng 595
Phyllocladus trichomanoides D. Don var. tricho­ Picea aurantiaca Mast. 551, 553, 561, 562, 596
manoides 549, 550 Picea aurantiaca Mast. var. retroflexa (Mast.)
1092 Picea A. Dietr. 17, 25, 38, 49, 50, 62–1112, 70, 262, C. T. Kuan & L. J. Zhou 595
263, 403, 404, 436, 444, 448, 459, 463, 468, 495, Picea austromandshurica Silba 574
551, 552, 553, 554, 560, 570, 590, 592, 633, 666, Picea balfouriana Rehd. & E. H. Wilson 579
717, 765, 932, 933, 981, 984, 1014, 1015, 1061 Picea bicolor (Maxim.) Mayr 557
Picea abies (L.) H. Karst. 25, 60, 65, 482, 551, 553, Picea bicolor (Maxim.) Mayr var. acicularis (Maxim.
554, 555, 556, 560, 570, 590, 591, 649, 732, 765, ex Beissn.) Shiras. 558
778, 1061 Picea bicolor (Maxim.) Mayr var. reflexa Shiras.  558
Picea abies (L.) H. Karst. var. abies 553, 555, 556 Picea brachytyla (Franch.) E. Pritz. 82, 551, 554,
Picea abies (L.) H. Karst. var. acuminata (Beck) 562, 563, 578, 955, 1061
Dallim. & A. B. Jacks. 555, 556 Picea brachytyla (Franch.) E. Pritz. var. ascendens
Picea abies (L.) H. Karst. var. alpestris (Brügger) (Patschke) Silba 563
P. A. Schmidt 556 Picea brachytyla (Franch.) E. Pritz. var. brachy­
Picea abies (L.) H. Karst. subsp. obovata (Ledeb.) tyla 74, 563
Hultén 555, 589 Picea brachytyla (Franch.) E. Pritz. var. complana­
Picea abies (L.) H. Karst. var. obovata (Ledeb.) ta (Mast.) W. C. Cheng ex Rehd. 115, 563, 934
Lindq. 589 Picea brachytyla (Franch.) E. Pritz. var. pachyclada
Picea acicularis Maxim. ex Beissn. 558 (Patschke) Silba 563
Picea ajanensis Fisch. ex Carrière 574 Picea brachytyla (Franch.) E. Pritz. var. rhombi­
Picea albertiana S. Br. 572 squamea Stapf 563
Picea albertiana S. Br. var. densata (L. H. Bailey) Picea breweriana S. Watson 277, 551, 554, 564,
W. L. Strong & Hills 572 565, 1030
Picea albertiana S. Br. subsp. ogilviei W. L. Strong & Picea chihuahuana Martínez 78, 551, 553, 565,
Hills 572 566, 583, 584, 626
Picea albertiana S. Br. var. porsildii (Raup) W. L. Picea complanata Mast. 563
Strong & Hills 572 Picea concolor Gordon 72
Picea alcoquiana (Veitch ex Lindl.) Carrière 551, Picea crassifolia Kom. 441, 551, 553, 566, 567
553, 556, 557, 559, 585 Picea engelmannii Parry ex Engelm. 97, 98, 113,
Picea alcoquiana (Veitch ex Lindl.) Carrière var. 401, 461, 491, 493, 552, 567, 568, 572, 609, 615,
acicularis (Maxim. ex Beissn.) Fitschen 558, 621, 654, 667, 676, 701, 715, 727, 761, 772, 932,
559, 577 993, 1017
Picea alcoquiana (Veitch ex Lindl.) Carrière var. Picea engelmannii Parry ex Engelm. subsp. engel­
alcoquiana 553, 557, 559 mannii 554, 568
Picea alcoquiana (Veitch ex Lindl.) Carrière var. Picea engelmannii Parry ex Engelm. var. glabra
reflexa (Shiras.) Fitschen 553, 558, 559 Goodman 568
Picea alpestris (Brügger) Stein 556 Picea engelmannii Parry ex Engelm. subsp. mexica­
Picea amabilis Douglas ex Loudon 61 na (Martínez) P. A. Schmidt 125, 126, 554, 568
Picea ascendens Patschke 563 Picea engelmannii Parry ex Engelm. var. mexicana
Picea asperata Mast. 82, 114, 122, 425, 551, 558, (Martínez) Silba 568
560, 561, 562, 567, 589, 595, 596, 605 Picea excelsa (Lam.) Link var. acuminata Beck 556
Picea asperata Mast. var. asperata 553, 560, 596 Picea farreri C. N. Page & Rushforth 551, 554, 569,
570, 1060
Picea ×fennica (Regel) Kom. 551, 555, 570, 571, Picea koraiensis Nakai var. pungsanensis (Uyeki ex
590 Nakai) Farjon 576
Picea ×fennica (Regel) Kom. subsp. uralensis (Tepl.) Picea koyamae Shiras. 551, 552, 576, 577, 585
P. A. Schmidt 570 Picea likiangensis (Franch.) E. Pritz. 74, 79, 84,
Picea fortunei A. Murray bis 475 445, 552, 554, 563, 578, 580, 581, 594, 595
Picea glauca (Moench) Voss 401, 490, 551, 552, Picea likiangensis (Franch.) E. Pritz. subsp. balfouri­
568, 571, 574, 581, 582, 597, 639, 654, 993, 1009, ana (Rehd. & E. H. Wilson) Rushforth 580
1017 Picea likiangensis (Franch.) E. Pritz. var. balfouriana
Picea glauca (Moench) Voss subsp. albertiana (Rehd. & E. H. Wilson) Hillier 579
(S. Br.) P. A. Schmidt 572 Picea likiangensis (Franch.) E. Pritz. var. bhutanica
Picea glauca (Moench) Voss var. albertiana (S. Br.) Silba 578 1093
Sarg. 571, 572 Picea likiangensis (Franch.) E. Pritz. var. forrestii
Picea glauca (Moench) Voss var. densata L. H. Bai- Silba 579
ley 572 Picea likiangensis (Franch.) E. Pritz. var. hirtella
Picea glauca (Moench) Voss subsp. engelmannii (Rehd. & E. H. Wilson) W. C. Cheng 579
(Engelm.) T. M. C. Taylor 568 Picea likiangensis (Franch.) E. Pritz. var. likiangen­
Picea glauca (Moench) Voss var. engelmannii sis 578, 594, 626
(Engelm.) Boivin 568 Picea likiangensis (Franch.) E. Pritz. var. linzhiensis
Picea glauca (Moench) Voss var. glauca 572 W. C. Cheng & L. K. Fu 580
Picea glauca (Moench) Voss var. porsildii Raup   572 Picea likiangensis (Franch.) E. Pritz. var. montigena
Picea glehnii (F. Schmidt) Mast. 117, 551, 553, 572, (Mast.) W. C. Cheng 579
573, 978 Picea likiangensis (Franch.) E. Pritz. var. purpurea
Picea hirtella Rehd. & E. H. Wilson 579 (Mast.) Dallim. & A. B. Jacks. 594
Picea hondoensis Mayr 575 Picea likiangensis (Franch.) E. Pritz. var. rubescens
Picea intercedens Nakai 576 Rehd. & E. H. Wilson 114, 122, 562, 579, 596
Picea jezoensis (Siebold & Zucc.) Carrière 96, Picea linzhiensis (W. C. Cheng & L. K. Fu) Rush-
105, 117, 484, 515, 552, 573, 574, 576, 978, 990, forth 122, 487, 552, 554, 580
994, 1013 Picea lowiana Gordon 72
Picea jezoensis (Siebold & Zucc.) Carrière var. Picea ×lutzii Little 552, 581
ajanensis (Fisch. ex Carrière) W. C. Cheng & Picea mariana (Mill.) Britton et al. 421, 490, 551,
L. K. Fu 574 554, 574, 581, 582, 597, 639, 991
Picea jezoensis (Siebold & Zucc.) Carrière subsp. Picea martinezii T. F. Patt. 551, 553, 583
hondoensis (Mayr) P. A. Schmidt 83, 94, 123, Picea maximowiczii Regel ex Mast. 551, 552, 577,
489, 554, 557, 574, 575 584
Picea jezoensis (Siebold & Zucc.) Carrière var. hon­ Picea maximowiczii Regel ex Mast. var. maximow­
doensis (Mayr) Rehd. 100, 575 iczii 585
Picea jezoensis (Siebold & Zucc.) Carrière subsp. Picea maximowiczii Regel ex Mast. var. senanensis
jezoensis 554 Hayashi 585
Picea jezoensis (Siebold & Zucc.) Carrière subsp. Picea mexicana Martínez 568
jezoensis var. jezoensis 574 Picea meyeri Rehd. & E. H. Wilson 551, 553, 585,
Picea jezoensis (Siebold & Zucc.) Carrière subsp. 586, 605
jezoensis var. komarovii (V. N. Vassil.) W. C. Picea meyeri Rehd. & E. H. Wilson var. mongolica
Cheng & L. K. Fu 575 H. Q. Wu 585
Picea jezoensis (Siebold & Zucc.) Carrière f. kam­ Picea meyeri Rehd. & E. H. Wilson f. pyramidalis
tchatkensis (Lacass.) S. L. Tung & Y. L. Chou 574 (H. W. Jen & C. G. Bai) L. K. Fu & Nan Li 585
Picea kamtchatkensis Lacass. 574 Picea meyeri Rehd. & E. H. Wilson var. pyramidalis
Picea komarovii V. N. Vassil. 575 H. W. Jen & C. G. Bai 585
Picea koraiensis Nakai 575, 576, 990 Picea montigena Mast. 579
Picea koraiensis Nakai var. intercedens (Nakai) Picea morinda Link subsp. tianschanica (Rupr.)
Y. L. Chou 576 Berezin 599
Picea koraiensis Nakai var. koraiensis 576
 Picea morrisonicola Hayata 95, 551, 552, 586, 587, Picea vulgaris Link var. uralensis Tepl. 570
609, 618, 620, 626, 674, 716, 729, 955 Picea watsoniana Mast. 604
Picea neoveitchii Mast. 82, 551, 552, 587, 588, 589, Picea wilsonii Mast. 551, 552, 586, 595, 604, 605,
595 628
Picea notabilis (Rehd. & E. H. Wilson) Lacass. 560 Picea wilsonii Mast. var. shanxiensis Silba 604
Picea obovata Ledeb. 93, 105, 120, 457, 481, 484, Picea wilsonii Mast. var. watsoniana (Mast.)
497, 551, 553, 555, 570, 576, 589, 590, 759, 1061 Silba 604
Picea obovata Ledeb. var. fennica (Regel) A. Hen- Pilgerodendron Florin 36, 46, 48, 370, 606
ry 570 Pilgerodendron uviferum (D. Don) Florin 370,
Picea omorika (Pančić) Purk. 551, 554, 590, 591 499, 606, 628, 872
1094 Picea orientalis (L.) Peterm. 107, 551, 553, 592, 627, Pinaceae Spreng. ex F. Rudolphi 16, 18, 22, 25, 26,
794 33, 37, 38, 43, 49, 55, 60, 251, 252, 254, 267, 472,
Picea pachyclada Patschke 563 475, 480, 533, 551, 576, 581, 595, 608, 684, 718,
Picea polita (Siebold & Zucc.) Carrière 602 924, 929, 944, 948, 956, 969, 1007, 1033, 1046,
Picea ponderosa (Rehd. & E. H. Wilson) Lacass.  561 1050, 1054, 1055, 1056, 1057, 1058, 1059, 1060,
Picea prostrata Isakov 598 1061, 1062, 1063
Picea pungens Engelm. 552, 554, 593, 609, 611, 752, Pinea Wolf 608
753 Pinus L. 13, 17, 24, 25, 27, 33, 37, 49, 50, 55, 73, 97,
Picea pungsanensis Uyeki ex Nakai 576 99, 113, 125, 190, 221, 249, 267, 277, 282, 295, 403,
Picea purpurea Mast. 82, 114, 552, 554, 589, 594, 412, 415, 418, 422, 431, 436, 448, 458, 464, 469,
595, 605 473, 475, 482, 489, 554, 562, 568, 574, 583, 590,
Picea purpurea Mast. var. balfouriana (Rehd. & 608, 609, 610, 612, 613, 614, 615, 616, 617, 634,
E. H. Wilson) Silba 579 653, 687, 695, 720, 725, 768, 795, 796, 858, 925,
Picea purpurea Mast. var. hirtella (Rehd. & E. H. 933, 946, 957, 1054
Wilson) Silba 579 Pinus abies L. 551, 556
Picea retroflexa Mast. 551, 553, 561, 595, 596 Pinus abies L. var. fennica Regel 570
Picea rubens Sarg. 87, 402, 551, 554, 596, 597, 798, Pinus albicaulis Engelm. 17, 428, 491, 609, 617,
799, 890, 1009 618, 619, 620, 621, 628, 638, 676, 715, 732, 932,
Picea schrenkiana Fisch. & C. A. Mey. 403, 411, 1029
442, 462, 551, 553, 597, 1068, 1069 Pinus amamiana Koidz. 609, 617, 619, 620
Picea schrenkiana Fisch. & C. A. Mey. subsp. Pinus apulcensis Lindl. 745
schrenkiana 598 Pinus araucana Molina 185, 191
Picea schrenkiana Fisch. & C. A. Mey. subsp. tian­ Pinus aristata Engelm. 98, 609, 616, 620, 621,
schanica (Rupr.) Bykov 598, 599, 627 628, 676, 703
Picea schrenkiana Fisch. & C. A. Mey. var. tian­ Pinus aristata Engelm. subsp. longaeva (D. K. Bai-
schanica (Rupr.) W. C. Cheng & S. H. Fu 599 ley) E. Murray 702
Picea shirasawae Hayashi 558 Pinus aristata Engelm. var. longaeva (D. K. Bailey)
Picea sitchensis (Bong.) Carrière 17, 61, 88, 113, Little 702
277, 552, 554, 574, 581, 591, 599, 627, 715, 993, Pinus arizonica Engelm. 292, 609, 615, 621, 622,
999, 1016, 1017, 1029 623, 651, 667, 701, 705, 739, 761, 772
Picea smithiana (Wall.) Boiss. 110, 121, 258, 551, Pinus arizonica Engelm. var. arizonica 624
552, 600, 601, 627, 628 Pinus arizonica Engelm. var. cooperi (C. E. Blanco)
Picea smithiana (Wall.) Boiss. var. nepalensis Fran- Farjon 624
co 600 Pinus arizonica Engelm. var. stormiae Martínez
Picea spinulosa (Griff.) Beissn. 77, 445, 487, 551, 624, 680, 751
554, 580, 581, 601 Pinus armandii Franch. 115, 569, 581, 587, 609, 617,
Picea spinulosa (Griff.) Beissn. var. yatungensis 620, 628, 633, 634, 662, 783, 794, 1069
Silba 601 Pinus armandii Franch. var. amamiana (Koidz.)
Picea tianschanica Rupr. 599 Hatus. 619, 620
Picea torano (Siebold ex K. Koch) Koehne 551, Pinus armandii Franch. var. armandii 628, 633
552, 602, 603, 604
Pinus armandii Franch. var. dabeshanensis Pinus californiarum D. K. Bailey subsp. fallax
(W. C. Cheng & Y. W. Law) Silba 634 (­Little) D. K. Bailey 711
Pinus armandii Franch. subsp. mastersiana (Haya- Pinus canadensis L. 1008
ta) Businský 634 Pinus canariensis C. Sm. 609, 612, 644, 645, 646
Pinus armandii Franch. var. mastersiana (Hayata) Pinus caribaea Morelet 17, 609, 610, 646, 647,
Hayata 95, 634 726, 727, 777, 843
Pinus attenuata Lemmon 304, 305, 309, 312, 393, Pinus caribaea Morelet var. bahamensis (Griseb.)
609, 611, 628, 634, 749 W. H. Barrett & Golfari 388, 648
Pinus ×attenuradiata Stockwell & Righter 635 Pinus caribaea Morelet var. caribaea 388, 647,
Pinus ayacahuite Ehrenb. ex Schltdl. 17, 89, 91, 777
308, 566, 609, 618, 635, 636, 666, 668, 705, 710, Pinus caribaea Morelet var. hondurensis (Sénécl.) 1095
731, 762, 771 W. H. Barrett & Golfari 647, 648
Pinus ayacahuite Ehrenb. ex Schltdl. var. ayaca­ Pinus catarinae Rob.-Pass. 750
huite 636 Pinus ceciliae Llorens & L. Llorens 681
Pinus ayacahuite Ehrenb. ex Schltdl. var. novo­ Pinus cembra L. 17, 555, 608, 609, 617, 619, 620,
galiciana Carvajal 761 629, 649, 650, 730, 732, 760
Pinus ayacahuite Ehrenb. ex Schltdl. subsp. strobi­ Pinus cembra L. var. pumila Pall. 745
formis (Engelm.) E. Murray 761 Pinus cembra L. subsp. sibirica (Du Tour) Kry-
Pinus ayacahuite Ehrenb. ex Schltdl. var. veitchii lov 758
(Roezl) Shaw 628, 636, 637 Pinus cembra L. var. sibirica (Du Tour) G. Don   758
Pinus bahamensis Griseb. 648 Pinus cembroides Zucc. 27, 292, 385, 391, 405, 406,
Pinus balfouriana Balf. 609, 616, 629, 637, 638, 415, 430, 435, 458, 609, 616, 650, 651, 665, 673,
703, 715 680, 701, 705, 721, 737, 744, 751
Pinus balfouriana Balf. subsp. aristata (Engelm.) Pinus cembroides Zucc. subsp. cembroides Little
Engelm. 620 var. bicolor Little 651
Pinus balfouriana Balf. var. aristata (Engelm.) Pinus cembroides Zucc. subsp. cembroides var.
Engelm. 620 cembroides 629, 651
Pinus balfouriana Balf. subsp. austrina Bruijn & Pinus cembroides Zucc. subsp. edulis (Engelm.)
J. Mastrog. 637, 638 E. Murray 669
Pinus balfouriana Balf. var. austrina (Bruijn & Pinus cembroides Zucc. var. edulis (Engelm.)
J. Mastrog.) Silba 637 Voss 669
Pinus balfouriana Balf. subsp. longaeva (D. K. Bai- Pinus cembroides Zucc. var. juarezensis (Lanner)
ley) E. Murray 702 Silba 747
Pinus balsamea L. 63 Pinus cembroides Zucc. subsp. lagunae (Rob.-Pass.)
Pinus banksiana Lamb. 17, 62, 401, 421, 490, 582, D. K. Bailey 652
609, 612, 639, 655, 752, 991 Pinus cembroides Zucc. var. lagunae Rob.-Pass.   652
Pinus bhutanica Grierson et al. 609, 617, 640 Pinus cembroides Zucc. subsp. monophylla (Torr. &
Pinus bracteata D. Don 66 Frém.) E. Murray 711
Pinus brutia Ten. 259, 314, 409, 412, 416, 431, 432, Pinus cembroides Zucc. subsp. orizabensis D. K.
434, 609, 610, 613, 641, 642, 643, 682, 722 Bailey 630, 652
Pinus brutia Ten. var. brutia 642 Pinus cembroides Zucc. var. orizabensis (D. K. Bai-
Pinus brutia Ten. var. densifolia F. Yaltirik & ley) Silba 652
M. Boydak 642 Pinus cembroides Zucc. var. quadrifolia (Parl. ex
Pinus brutia Ten. var. eldarica (Medw.) Silba 642 Sudw.) Silba 747
Pinus brutia Ten. var. pendulifolia Frankis 642 Pinus cembroides Zucc. var. remota Little 750
Pinus brutia Ten. var. pityusa (Steven) Silba 643 Pinus chiapensis (Martínez) Andresen 618, 763,
Pinus brutia Ten. var. stankewiczii (Sukaczev) 1053
Frankis 643 Pinus chihuahuana Engelm. 702
Pinus bungeana Zucc. ex Endl. 27, 473, 609, 616, Pinus cilicica Antoine & Kotschy 72
629, 643, 678, 760, 1069 Pinus clausa (Chapm. ex Engelm.) Sarg. 609, 612,
Pinus californiarum D. K. Bailey 711 652, 653
 Pinus clausa (Chapm. ex Engelm.) Sarg. subsp. im­ Pinus densiflora Siebold & Zucc. f. sylvestriformis
muginata (D. B. Ward) E. Murray 652 Taken. 663
Pinus clausa (Chapm. ex Engelm.) Sarg. var. im­ Pinus densiflora Siebold & Zucc. var. sylvestriformis
muginata D. B. Ward 652, 653 (Taken.) Q. L. Wang 663
Pinus clusiana Clemente 724 Pinus densiflora Siebold & Zucc. var. ussuriensis
Pinus contorta Douglas ex Loudon 304, 428, 461, T. N. Liou & Q. L. Wang 664
582, 593, 609, 612, 619, 654, 655, 676, 690, 715, Pinus densiflora Siebold & Zucc. var. zhangwuensis
932, 993, 1029, 1030 S. J. Zhang et al. 663, 664
Pinus contorta Douglas ex Loudon var. contor­ Pinus ×densithunbergii Uyeki 609, 662
ta 655 Pinus densi-thunbergii Uyeki 662
1096 Pinus contorta Douglas ex Loudon subsp. latifolia Pinus deodara Lamb. 256
(Engelm.) Critchf. 655 Pinus devoniana Lindl. 89, 609, 615, 630, 664,
Pinus contorta Douglas ex Loudon var. latifolia 665, 666, 710, 727, 741, 742
Engelm. 493, 655 Pinus discolor D. K. Bailey & Hawksw. 651
Pinus contorta Douglas ex Loudon subsp. mur­ Pinus divaricata (Aiton) Dum.-Cours. var. latifolia
rayana (Balf.) Engelm. 655 (Engelm.) Boivin 655
Pinus contorta Douglas ex Loudon var. murrayana Pinus donnell-smithii Mast. 683
(Balf.) Engelm. 630, 654, 655, 696 Pinus douglasiana Martínez 609, 615, 666, 667,
Pinus cooperi C. E. Blanco 624 689, 701, 705, 710, 727, 742
Pinus coulteri D. Don 66, 247, 292, 609, 615, 630, Pinus douglasiana Martínez var. martinezii
655, 690, 757, 931 (E. Larsen) Silba 667
Pinus crassicorticea Y. C. Zhong & K. X. Huang   707 Pinus douglasiana Martínez var. maximinoi
Pinus cubensis Griseb. 609, 610, 656, 657 (H. E. Moore) Silba 709
Pinus culminicola Andresen & Beaman 17, 609, Pinus douglasii Sabine ex D. Don 929
616, 630, 657, 658 Pinus dumosa D. Don 1013
Pinus culminicola Andresen & Beaman var. discolor Pinus durangensis Martínez 78, 609, 615, 630,
(D. K. Bailey & Hawksw.) Silba 651 667, 668, 701, 761, 772
Pinus culminicola Andresen & Beaman var. johan­ Pinus echinata Mill. 470, 609, 611, 668, 669, 679,
nis (Rob.-Pass.) Silba 651 728, 753, 779
Pinus cupressoides Molina 370 Pinus edulis Engelm. 27, 292, 425, 430, 461, 608,
Pinus dabeshanensis W. C. Cheng & Y. W. Law 634 609, 616, 669, 670
Pinus dalatensis Ferré 609, 618, 659, 660, 695, Pinus edulis Engelm. var. fallax Little 711
729, 781, 1054 Pinus eldarica Medw. 642
Pinus dalatensis Ferré subsp. dalatensis var. Pinus elliottii Engelm. 26, 470, 609, 612, 670, 671,
bidoup­ensis Businský 661 679, 728, 758, 980
Pinus dalatensis Ferré subsp. dalatensis var. dalat­ Pinus elliottii Engelm. subsp. densa (Little &
ensis 661 K. W. Dorman) E. Murray 672
Pinus dalatensis Ferré subsp. procera Businský 661 Pinus elliottii Engelm. var. densa Little &
Pinus dalmatica Vis. 723 K. W. Dorman 672
Pinus dammara Lamb. 143, 150, 151 Pinus elliottii Engelm. var. elliottii 671
Pinus densa (Little & K. W. Dorman) Silba 672 Pinus engelmannii Carrière 623, 630, 651, 672, 673
Pinus densa (Little & K. W. Dorman) Silba var. aus­ Pinus engelmannii Carrière var. blancoi (Martínez)
trokeysensis Silba 672 Martínez 672
Pinus densata Mast. 300, 374, 609, 614, 661, 662, Pinus eremitana Businský 673, 674
782 Pinus escarena Risso 735
Pinus densata Mast. var. pygmaea J. R. Xue 783 Pinus estevezii (Martínez) J. P. Perry 744
Pinus densiflora Siebold & Zucc. 83, 94, 280, 446, Pinus ×fallax (Little) Businský 711
489, 584, 604, 609, 614, 662, 663, 664, 769, 1019 Pinus fenzeliana Hand.-Mazz. 173, 253, 519, 535,
Pinus densiflora Siebold & Zucc. var. densiflora 664 609, 618, 620, 634, 673, 674, 729, 781, 975
Pinus densiflora Siebold & Zucc. var. funebris Pinus fenzeliana Hand.-Mazz. var. annamiensis
(Kom.) T. N. Liou & Q. L. Wang ex Silba 663 Silba 673
Pinus fenzeliana Hand.-Mazz. var. dabeshanensis Pinus hwangshanensis W. Y. Hsia 609, 614, 687,
(W. C. Cheng & Y. W. Law) L. K. Fu & Nan 688, 769
Li 634 Pinus inops Aiton var. clausa Chapm. ex
Pinus flexilis E. James 98, 401, 461, 491, 609, 617, Engelm. 652
636, 674, 675, 676, 677, 703, 715, 762, 932 Pinus insignis Douglas ex Loudon var. binata
Pinus flexilis E. James var. flexilis 676 Engelm. 749
Pinus flexilis E. James var. macrocarpa Engelm.  676 Pinus insularis Endl. 692, 782, 800, 801
Pinus flexilis E. James subsp. reflexa (Engelm.) Pinus insularis Endl. var. khasyana (Griff.) Silba  692
E. Murray 677 Pinus insularis Endl. var. langbianensis (A. Chev.)
Pinus flexilis E. James var. reflexa Engelm. 676, Silba 692
677 Pinus insularis Endl. var. tenuifolia (W. C. Cheng & 1097
Pinus formosana Hayata 716 Y. W. Law) Silba 783
Pinus fragilissima Businský 769, 770 Pinus insularis Endl. var. yunnanensis (Franch.)
Pinus fraseri Pursh 86 Silba 783
Pinus funebris Kom. 663, 664 Pinus jaliscana Pérez de la Rosa 609, 611, 688,
Pinus gerardiana Wall. ex D. Don 27, 414, 609, 689
616, 677, 678, 760 Pinus jeffreyi Balf. 247, 292, 295, 428, 609, 615,
Pinus glabra Walter 609, 610, 611, 678, 679, 1006 689, 690, 696, 712, 715, 739, 748, 931, 951
Pinus glauca Moench 572 Pinus jeffreyi Balf. var. baja-californica Silba 689
Pinus gordoniana Hartw. ex Gordon 714 Pinus johannis Rob.-Pass. 651
Pinus grandis Douglas ex D. Don 87 Pinus juarezensis Lanner 747
Pinus greggii Engelm. ex Parl. 609, 611, 679, 731 Pinus kaempferi Lamb. 488
Pinus greggii Engelm. ex Parl. var. australis Dona- Pinus kesiya Royle ex Gordon 17, 26, 609, 610,
hue & Lopez 680 614, 661, 691, 692, 711, 782, 783
Pinus greggii Engelm. ex Parl. var. greggii 680 Pinus kesiya Royle ex Gordon subsp. insularis
Pinus ×hakkodensis Makino 609, 681 (Endl.) D. Z. Li 692
Pinus halepensis Mill. 27, 431, 434, 609, 613, 642, Pinus kesiya Royle ex Gordon var. kesiya 692
681, 682, 722, 738 Pinus kesiya Royle ex Gordon var. langbianensis
Pinus halepensis Mill. subsp. brutia (Ten.) Holm- (A. Chev.) Gaussen ex N.-S. Bui 692, 782
boe 642 Pinus kesiya Royle ex Gordon subsp. yunnanensis
Pinus halepensis Mill. var. brutia (Ten.) A. Hen- (Franch.) Businský 783
ry 642 Pinus kochiana Klotzsch ex K. Koch 766
Pinus halepensis Mill. var. ceciliae (Llorens & L. Llo- Pinus koraiensis Siebold & Zucc. 27, 93, 96, 105,
rens) Roselló et al. 681 446, 457, 515, 551, 553, 609, 617, 692, 693, 978,
Pinus hamata (Steven) Sosn. 766 990
Pinus hartwegii Lindl. 89, 116, 308, 428, 464, 609, Pinus krempfii Lecomte 608, 609, 616, 659, 693,
615, 631, 658, 665, 683, 684, 761 694, 695
Pinus hartwegii Lindl. var. rudis (Endl.) Silba 683 Pinus krempfii Lecomte var. poilanei Lecomte 693
Pinus ×hayatana Businský 716 Pinus kwangtungensis Chun & Tsiang 169, 251,
Pinus heldreichii H. Christ. 609, 610, 612, 631, 253, 270, 519, 535, 673, 674, 729, 781, 975
684, 685, 722 Pinus kwangtungensis Chun & Tsiang var. variifolia
Pinus heldreichii H. Christ subsp. leucodermis Nan Li & Y. C. Zhong 673, 674
(­Antoine) E. Murray 684 Pinus lagunae (Rob.-Pass.) Passini 652
Pinus heldreichii H. Christ var. leucodermis (An­ Pinus lambertiana Douglas 17, 247, 292, 312, 564,
toine) Markgr. ex Fitschen 684, 685 609, 617, 690, 696, 697, 715, 951, 993
Pinus henryi Mast. 609, 614, 685, 686 Pinus lambertiana Douglas var. martirensis
Pinus herrerae Martínez 609, 611, 666, 686, 687, Silba 696
700, 756 Pinus lanceolata Lamb. 286, 288
Pinus hingganensis H. J. Zhang 758 Pinus langbianensis A. Chev. 692
Pinus ×holfordiana A. B. Jacks 636 Pinus laricina Du Roi 489
Pinus hondurensis Sénécl. 648 Pinus laricio Poir. 723
 Pinus laricio Poir. var. angustisquama Willk. 724 Pinus massoniana Lamb. var. shaxianensis
Pinus laricio Poir. subsp. calabrica (Loudon) Cesca D. X. Zhou 707
& Peruzzi 723 Pinus massoniana Lamb. var. wulingensis C. J. Qi &
Pinus laricio Poir. var. calabrica Loudon 723 Q. Z. Lin 685
Pinus laricio Poir. var. caramanica Loudon 724 Pinus mastersiana Hayata 634
Pinus laricio Poir. var. latisquama Willk. 724 Pinus maximartinezii Rzed. 17, 609, 617, 631, 708
Pinus larix L. 489, 497 Pinus maximinoi H. E. Moore 308, 609, 615, 665,
Pinus larix L. var. russica Endl. 496, 497 689, 709, 727, 731, 742, 771
Pinus lasiocarpa Hook. 98 Pinus merkusii Jungh. & de Vriese 609, 613, 699,
Pinus latteri Mason 609, 613, 631, 698, 699, 711 710, 711, 1054
1098 Pinus lawsonii Roezl ex Gordon 609, 612, 666, Pinus merkusii Jungh. & de Vriese subsp. latteri
699, 700, 701, 742 (Mason) D. Z. Li 698
Pinus lawsonii Roezl ex Gordon var. gracilis Debre- Pinus merkusii Jungh. & de Vriese var. latteri
czy & Rácz 699 (­Mason) Silba 698
Pinus leiophylla Schiede ex Schltdl. & Cham. 292, Pinus merkusii Jungh. & de Vriese subsp. ustulata
405, 609, 610, 666, 667, 673, 700, 705, 727, 772 Businský 710
Pinus leiophylla Schiede ex Schltdl. & Cham. subsp. Pinus mertensiana Bong. 1007, 1018
chihuahuana (Engelm.) E. Murray 702 Pinus mesogeensis Fieschi & Gaussen 734, 735
Pinus leiophylla Schiede ex Schltdl. & Cham. var. Pinus michoacana Martínez 664, 665
chihuahuana (Engelm.) Shaw 78, 651, 701, Pinus monophylla Torr. & Frém. 27, 292, 430, 609,
702, 708, 761 613, 616, 631, 632, 711, 712, 748
Pinus leiophylla Schiede ex Schltdl. & Cham. var. Pinus monophylla Torr. & Frém. var. californiarum
leiophylla 701 (D. K. Bailey) Silba 711
Pinus leucodermis Antoine 684, 685 Pinus monophylla Torr. & Frém. var. fallax (Little)
Pinus longaeva D. K. Bailey 17, 609, 616, 621, 631, Silba 711
638, 676, 702, 703, 712 Pinus montezumae Lamb. 89, 90, 116, 308, 385,
Pinus ×longipedunculata (Loock ex Martínez) 609, 615, 665, 668, 680, 683, 700, 701, 712, 713,
Businský 731 772
Pinus luchuensis Mayr 609, 614, 688, 703, 704, Pinus montezumae Lamb. var. gordoniana (Hartw.
769 ex Gordon) Silba 714
Pinus luchuensis Mayr subsp. hwangshanensis Pinus montezumae Lamb. var. mezambrana Carva-
(W. Y. Hsia) D. Z. Li 687 jal 714
Pinus luchuensis Mayr var. hwangshanensis Pinus montezumae Lamb. var. montezumae 714
(W. Y. Hsia) C. L. Wu 687 Pinus montezumae Lamb. var. rudis (Endl.)
Pinus lumholtzii B. L. Rob & Fernald 609, 610, Shaw 683
673, 704, 705, 741, 761 Pinus monticola Douglas ex D. Don 247, 428, 493,
Pinus lumholtzii B. L. Rob & Fernald var. micro­ 564, 609, 618, 636, 690, 696, 714, 715, 739, 780,
phylla Carvajal 701 993, 1029
Pinus luzmariae Pérez de la Rosa 609, 611, 705 Pinus morrisonicola Hayata 716
Pinus macrophylla Engelm. var. blancoi Martín- Pinus mugo Turra 403, 609, 613, 658, 716, 717, 722,
ez 672 752, 778, 1054
Pinus macvaughii Carvajal 688 Pinus mugo Turra subsp. mugo 718
Pinus maestrensis Bisse 656 Pinus mugo Turra subsp. rostrata (Antoine) E. Mur-
Pinus martinezii E. Larsen 667 ray 777
Pinus massoniana Lamb. 26, 289, 374, 473, 535, Pinus mugo Turra var. rostrata (Antoine)
609, 614, 664, 686, 706 Hoopes 777
Pinus massoniana Lamb. var. hainanensis W. C. Pinus mugo Turra subsp. rotundata (Link) Janch. &
Cheng & L. K. Fu 707 H. Neumayer 717, 718, 778
Pinus massoniana Lamb. var. henryi (Mast.) Pinus mugo Turra subsp. rotundata (Link) Janch.
C. L. Wu 685 & H. Neumayer var. pseudopumilio (Willk.)
Pinus massoniana Lamb. var. massoniana 707 P. Schmidt 718
Pinus mugo Turra subsp. uncinata (Ramond ex DC) Pinus nigra J. F. Arnold subsp. salzmannii (Dunal)
Domin 778 Franco 632, 724, 734
Pinus mukdensis Uyeki ex Nakai 767 Pinus nigra J. F. Arnold var. salzmannii (Dunal)
Pinus muricata D. Don 304, 609, 610, 632, 635, Laguna Lumbreras 724
718, 719 Pinus nigra J. F. Arnold var. yaltirikiana C. U. Alpte­
Pinus muricata D. Don var. borealis Axelrod ex Far- kin 724
jon 718 Pinus nordmanniana Steven 108
Pinus muricata D. Don var. cedrosensis J. T. How- Pinus novaemexicana P. Landry 676
ell 749 Pinus nubicola J. P. Perry 744
Pinus muricata D. Don var. stantonii Axelrod ex Pinus oaxacana Mirov 745
Farjon 718 Pinus oaxacana Mirov var. diversiformis Debreczy 1099
Pinus murrayana Balf. 655 & Rácz 745
Pinus ×neilreichiana Reichardt 609, 719 Pinus occidentalis Sw. 388, 609, 611, 725, 1055
Pinus nelsonii Shaw 435, 609, 615, 632, 651, 720, Pinus occidentalis Sw. var. baorucoensis Silba 725
721 Pinus occidentalis Sw. var. cubensis (Griseb.)
Pinus nigra J. F. Arnold 69, 72, 259, 409, 412, 416, Silba 656
431, 555, 591, 609, 612, 614, 664, 684, 685, 720, Pinus occidentalis Sw. var. maestrensis (Bisse)
721, 722, 723, 773, 1069 Silba 656
Pinus nigra J. F. Arnold var. angustisquama (Willk.) Pinus omorika Pančić 590
Laguna Lumbreras 724 Pinus oocarpa Schiede ex Schltdl. 418, 609, 611,
Pinus nigra J. F. Arnold subsp. calabrica (Loudon) 648, 665, 666, 668, 689, 700, 701, 705, 706, 710,
E. Murray 723 726, 727, 741, 742, 756, 771, 772, 843
Pinus nigra J. F. Arnold var. calabrica (Loudon) Pinus oocarpa Schiede ex Schltdl. var. macvaughii
C. K. Schneid. 723 (Carvajal) Silba 688
Pinus nigra J. F. Arnold subsp. caramanica (Lou- Pinus oocarpa Schiede ex Schltdl. var. manzanoi
don) Businský 724 Martínez 726
Pinus nigra J. F. Arnold var. caramanica (Loudon) Pinus oocarpa Schiede ex Schltdl. var. ochoterenae
Rehd. 724 Martínez 770
Pinus nigra J. F. Arnold subsp. clusiana (Clemente) Pinus oocarpa Schiede ex Schltdl. f. trifoliata Mar-
Rivas-Martínez 724 tínez 705
Pinus nigra J. F. Arnold var. columnaris-pendula Pinus oocarpa Schiede ex Schltdl. var. trifoliata
Boydak 724 Martínez 705
Pinus nigra J. F. Arnold subsp. croatica Lovric 723 Pinus orientalis L. 592
Pinus nigra J. F. Arnold subsp. dalmatica (Vis.) Pinus orizabensis (D. K. Bailey) D. K. Bailey &
Franco 723 Hawksw. 652
Pinus nigra J. F. Arnold var. dalmatica (Vis.) Busin- Pinus orthophylla Businský 673, 674
ský 723 Pinus pallasiana Lamb. 724
Pinus nigra J. F. Arnold subsp. laricio (Poir.) Pinus pallasiana Lamb. subsp. caramanica (Lou-
Maire 632, 722, 723 don) Chrtek & B. Slavik 724
Pinus nigra J. F. Arnold var. latisquama (Willk.) Pinus palustris Mill. 17, 26, 609, 611, 665, 727, 728,
Laguna Lumbreras 724 753, 980
Pinus nigra J. F. Arnold subsp. nigra 685, 720, 723 Pinus parviflora Siebold & Zucc. 83, 123, 280, 557,
Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.) 609, 617, 716, 728, 729, 782, 947, 994, 996, 1013,
Holmboe 432, 723, 724 1019
Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.) Pinus parviflora Siebold & Zucc. var. fenzeliana
Holmboe var. fastigiata Businský 724 (Hand.-Mazz.) C. L. Wu 673
Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.) Pinus parviflora Siebold & Zucc. var. morrisonicola
Holmboe var. pallasiana 724 (Hayata) C. L. Wu 716
Pinus nigra J. F. Arnold var. pyramidata A. Acatay Pinus parviflora Siebold & Zucc. var. parvi­flora 730
(nom. inval.) 724 Pinus parviflora Siebold & Zucc. subsp. pentaphylla
(Mayr) Businský 730
 Pinus parviflora Siebold & Zucc. var. pentaphylla Pinus ponderosa Douglas ex C. Lawson var. ponde­
(Mayr) A. Henry 681, 729, 730 rosa 739, 740, 785
Pinus patula Schiede ex Schltdl. & Cham. 92, 308, Pinus ponderosa Douglas ex C. Lawson subsp.
609, 610, 680, 700, 701, 710, 730, 731, 742, 772, ­scopulorum (Engelm.) E. Murray 740
1025 Pinus ponderosa Douglas ex C. Lawson var. scopu­
Pinus patula Schiede ex Schltdl. & Cham. var. jalis­ lorum Engelm. 739, 740
cana (Pérez de la Rosa) Silba 688 Pinus ponderosa Douglas ex C. Lawson var.
Pinus patula Schiede ex Schltdl. & Cham. var. ­stormiae (Martínez) Silba 624
longi­pedunculata Loock ex Martínez 731 Pinus ponderosa Douglas ex C. Lawson subsp.
Pinus patula Schiede ex Schltdl. & Cham. var. washoensis (Mason & Stockw.) E. Murray 740
1100 patula 632, 731 Pinus praetermissa Styles & McVaugh 609, 611,
Pinus patula Schiede ex Schltdl. & Cham. subsp. 740, 741
tecunumanii (Eguiluz & J. P. Perry) Styles 770 Pinus pringlei Shaw 609, 612, 700, 701, 741, 742
Pinus pentaphylla Mayr 729, 730 Pinus prokoraiensis Y. T. Zhao 692
Pinus pentaphylla Mayr var. hakkodensis (Makino) Pinus pseudostrobus Lindl. 89, 91, 92, 308, 566,
Kusaka 681 609, 615, 651, 665, 666, 673, 680, 700, 701, 710,
Pinus peuce Griseb. 609, 618, 722, 732 731, 741, 742, 743, 744, 756, 771
Pinus peuce var. vermiculata Christ 733 Pinus pseudostrobus Lindl. subsp. apulcensis
Pinus picea L. 55 (Lindl.) Stead 744
Pinus pinaster Aiton 26, 112, 609, 610, 612, 733, Pinus pseudostrobus Lindl. var. apulcensis (Lindl.)
734, 735 Martínez 744
Pinus pinaster Aiton subsp. acutisquama (Boiss.) Pinus pseudostrobus Lindl. var. apulcensis (Lindl.)
Rivas-Martínez 734 Shaw 745
Pinus pinaster Aiton var. acutisquama Boiss. 734 Pinus pseudostrobus Lindl. var. estevezii Martínez
Pinus pinaster Aiton subsp. atlantica Villar 734 744
Pinus pinaster Aiton subsp. escarena (Risso) Pinus pseudostrobus Lindl. var. laubenfelsii
K. Richt. 734, 735 Silba 744
Pinus pinaster Aiton var. mesogeensis (Fieschi & Pinus pseudostrobus Lindl. var. oaxacana (Mirov)
Gaussen) Silba 735 S. G. Harrison 745
Pinus pinaster Aiton subsp. pinaster 734, 785 Pinus pseudostrobus Lindl. var. pseudostrobus 744
Pinus pinaster Aiton subsp. renoui (Villar) Pinus pumila (Pall.) Regel 17, 100, 105, 117, 123,
Maire 734, 735 457, 484, 515, 574, 609, 617, 658, 681, 745, 990
Pinus pinaster Aiton var. renoui Villar 735 Pinus pungens Lamb. 746, 747, 785
Pinus pinceana Gordon 435, 609, 617, 651, 721, Pinus quadrifolia Parl. ex Sudw. 609, 616, 712,
735, 736 747, 748, 785
Pinus pindrow Royle ex D. Don 110 Pinus radiata D. Don 23, 230, 304, 311, 312, 609,
Pinus pinea L. 27, 431, 608, 609, 613, 722, 737, 738, 611, 635, 680, 719, 748, 749, 894
785 Pinus radiata D. Don subsp. binata (Engelm.)
Pinus pityusa Steven 643 E. Murray 749
Pinus pityusa Steven var. stankewiczii Sukac- Pinus radiata D. Don var. binata (Engelm.) Lem-
zev 643 mon 306, 749, 750
Pinus ponderosa Douglas ex C. Lawson 66, 247, Pinus radiata D. Don var. cedrosensis (J. T. Howell)
292, 295, 304, 305, 309, 401, 428, 461, 493, 564, Silba 749
609, 610, 614, 623, 624, 670, 690, 696, 699, 712, Pinus radiata D. Don var. radiata 749
715, 738, 739, 931, 932, 951, 993, 1058 Pinus reflexa (Engelm.) Engelm. 677
Pinus ponderosa Douglas ex C. Lawson subsp. ari­ Pinus religiosa Kunth 116
zonica (Engelm.) E. Murray 624 Pinus remota (Little) D. K. Bailey & Hawksw. 387,
Pinus ponderosa Douglas ex C. Lawson var. arizoni­ 609, 616, 721, 750, 751
ca (Engelm.) Shaw 624 Pinus renoui (Villar) Gaussen 735
Pinus ponderosa Douglas ex C. Lawson subsp. coul­ Pinus resinosa Aiton 609, 613, 639, 751, 752, 1009
teri (D. Don) E. Murray 655 Pinus ×rhaetica Brügger 609, 752
Pinus rigida Mill. 609, 611, 747, 752, 753, 758, 779 Pinus sylvestris L. var. mongolica Litv. 766
Pinus rotundata Link 718 Pinus sylvestris L. var. scotica Beissn. 765
Pinus roxburghii Sarg. 609, 612, 754, 755, 780 Pinus sylvestris L. var. sylvestriformis (Taken.)
Pinus rudis Endl. 683 W. C. Cheng & C. D. Chu 663
Pinus rzedowskii Madrigal & M. Caball. 609, 617, Pinus sylvestris L. var. sylvestris 764, 765, 786
755, 786 Pinus tabuliformis Carrière 289, 609, 614, 643,
Pinus sabiniana Douglas ex D. Don 292, 309, 312, 662, 686, 766, 767, 782, 783, 794
393, 609, 615, 756, 757 Pinus tabuliformis Carrière var. brevifolia S. Y. Wang
Pinus salzmannii Dunal 724 & C. L. Chang 767
Pinus ×schwerinii Fitschen 780 Pinus tabuliformis Carrière var. densata (Mast.)
Pinus scopulorum (Engelm.) Lemmon 740 Rehd. 661 1101
Pinus serotina Michx. 609, 611, 671, 757, 758 Pinus tabuliformis Carrière subsp. henryi (Mast.)
Pinus sibirica Du Tour 27, 105, 120, 442, 497, 576, Businský 685
609, 617, 758, 759, 990 Pinus tabuliformis Carrière var. henryi (Mast.)
Pinus sibirica Du Tour var. hingganensis (H. J. C. T. Kuan 685
Zhang) Silba 758 Pinus tabuliformis Carrière subsp. mukdensis
Pinus sitchensis Bong. 599 (Uyeki ex Nakai) Businský 767
Pinus smithiana Wall. 600 Pinus tabuliformis Carrière var. mukdensis (Uyeki
Pinus sosnowskyi Nakai 766 ex Nakai) Uyeki 767
Pinus spectabilis D. Don 120 Pinus tabuliformis Carrière var. pygmaea (J. R. Xue)
Pinus squamata X. W. Li 609, 616, 760 Silba 783
Pinus strobiformis Engelm. 78, 566, 609, 618, 623, Pinus tabuliformis Carrière var. tabuliformis 767
677, 761, 762 Pinus tabuliformis Carrière var. umbraculifera
Pinus strobiformis Engelm. var. carvajalii Silba 761 T. N. Liou & Q. L. Wang 767
Pinus strobiformis Engelm. var. potosiensis Pinus tabuliformis Carrière var. yunnanensis Dal-
Silba 677 lim. & A. B. Jacks. 783
Pinus strobiformis Engelm. subsp. veitchii (Roezl) Pinus taeda L. 609, 611, 669, 671, 679, 728, 758,
Frankis 637 768, 779, 1006
Pinus strobus L. 62, 597, 608, 609, 618, 636, 715, Pinus taiwanensis Hayata 287, 609, 614, 688, 769,
742, 752, 761, 762, 763, 780, 991, 1009 770
Pinus strobus L. subsp. chiapensis (Martínez) Pinus taiwanensis Hayata var. damingshanensis
E. Murray 763 W. C. Cheng & L. K. Fu 687, 688
Pinus strobus L. var. chiapensis Martínez 715, 763, Pinus taiwanensis Hayata var. fragilissima (Busin-
771, 981 ský) Farjon 770
Pinus strobus L. subsp. monticola (Douglas ex Pinus taiwanensis Hayata var. taiwanensis 770,
D. Don) E. Murray 714 786
Pinus strobus L. var. monticola (Douglas ex D. Don) Pinus tecunumanii Eguiluz & J. P. Perry 609, 611,
Nuttall 714 648, 710, 727, 770, 771
Pinus strobus L. var. strobus 763 Pinus tenuifolia Benth. 709
Pinus stylesii Frankis ex Businský 677 Pinus teocote Schiede ex Schltdl. & Cham. 27, 92,
Pinus sylvestris L. 17, 25, 60, 107, 401, 481, 483, 405, 609, 612, 668, 680, 700, 701, 705, 772
484, 497, 555, 570, 590, 608, 609, 614, 662, 664, Pinus teocote Schiede ex Schltdl. & Cham. var. her­
720, 721, 722, 746, 752, 759, 764, 765, 778, 801, rerae (Martínez) Silba 686
856, 1054, 1069 Pinus thunbergiana Franco 773
Pinus sylvestris L. subsp. hamata (Steven) Fomin Pinus thunbergii Parl. 609, 613, 663, 664, 773
766 Pinus torreyana Parry ex Carrière 609, 615, 774,
Pinus sylvestris L. var. hamata Steven 766 775
Pinus sylvestris L. subsp. kochiana (Klotzsch ex Pinus torreyana Parry ex Carrière subsp. insularis
K. Koch) Eliçin 766 J. R. Haller 776
Pinus sylvestris L. var. manguiensis S. Y. Li & Pinus torreyana Parry ex Carrière var. insularis
Adair 766 (J. R. Haller) Silba 776
 Pinus torreyana Parry ex Carrière subsp. torreya­ Platycladus orientalis (L.) Franco 396, 786, 793,
na 775, 776 794, 1057, 1059, 1069
Pinus tropicalis Morelet 388, 609, 614, 776, 777 Platycladus stricta Spach 793
Pinus uliginosa Neuman ex Wimm. 718 Platycyparis A. V. Bobrov & Melikyan 290
Pinus uncinata Ramond ex DC. 609, 613, 718, Platycyparis funebris (Endl.) A. V. Bobrov & Me-
777, 778 likyan 290, 302
Pinus uncinata Ramond ex DC var. ancestralis Podocarpaceae Endl. 11, 12, 15, 16, 17, 18, 19, 22,
Businský 777 24, 26, 33, 39, 43, 48, 49, 51, 129, 136, 204, 208,
Pinus uncinata Ramond ex DC var. pseudopumilio 251, 317, 337, 353, 360, 361, 363, 367, 377, 478, 498,
Willk. 718 510, 511, 516, 518, 539, 541, 546, 795, 850, 852,
1102 Pinus uncinata Ramond ex DC var. rostrata An­ 884, 913, 916, 921, 922, 937, 938, 944, 952, 1038,
toine 777 1047, 1053, 1054, 1056, 1058, 1059, 1060, 1062
Pinus uncinata Ramond ex DC subsp. uliginosa Podocarpus L’Hér. ex Pers. 13, 17, 18, 22, 26, 39, 51,
(Neumann ex Wimm.) Businský 718 136, 138, 139, 140, 148, 202, 222, 319, 320, 323,
Pinus uyematsui Hayata 716 324, 337, 347, 475, 504, 524, 537, 539, 547, 795,
Pinus veitchii Roezl 637 796, 797, 798, 799, 800, 801, 802, 804, 812, 826,
Pinus virginiana Mill. 469, 609, 612, 747, 752, 753, 828, 843, 844, 847, 862, 868, 881, 885, 938, 943,
778, 779, 786 952
Pinus wallichiana A. B. Jacks. 110, 121, 258, 414, Podocarpus acuminatus de Laub. 803, 805
442, 445, 462, 487, 563, 601, 602, 609, 617, 636, Podocarpus acutifolius Kirk 787, 801, 804, 806
640, 661, 754, 779, 780, 786 Podocarpus affinis Seem. 801, 807
Pinus wallichiana A. B. Jacks. subsp. bhutanica Podocarpus allenii Standl. 842
(­Grierson et al.) Businský 640 Podocarpus alpinus R. Br. ex Hook. f. 851
Pinus wallichiana A. B. Jacks. var. manangensis Podocarpus alpinus R. Br. ex Hook. f. var. arbores­
H. Ohba & M. Suzuki 780 cens Brongn. & Gris 840
Pinus wallichiana A. B. Jacks. var. parva K. C. Podocarpus alpinus R. Br. ex Hook. f. var. caespito­
­Sahni 781 sus Pancher ex Brongn. & Gris 840
Pinus wallichiana A. B. Jacks. var. wallichiana 780 Podocarpus alpinus R. Br. ex Hook. f. var. lawrencei
Pinus wangii Hu & W. C. Cheng 609, 618, 674, (Hook. f.) Hook. f. 851
729, 781, 782 Podocarpus amarus Blume 952
Pinus wangii Hu & W. C. Cheng subsp. kwangtung­ Podocarpus andinus Poepp. ex Endl. 913, 914
ensis (Chun & Tsiang) Businský 673 Podocarpus angustifolius Griseb. 802, 807, 808,
Pinus wangii Hu & W. C. Cheng var. kwangtungen­ 832, 876
sis (Chun & Tsiang) Silba 673 Podocarpus angustifolius Griseb. var. aristulatus
Pinus wangii Hu & W. C. Cheng subsp. variifolia (Parl.) Staszk. 807
(Nan Li & Y. C. Zhong) Businský 673 Podocarpus angustifolius Griseb. subsp. buchii
Pinus washoensis Mason & Stockw. 740 (Urb.) Staszk. 818
Pinus yecorensis Debreczy & Rácz 744 Podocarpus angustifolius Griseb. subsp. ­buchii
Pinus yecorensis Debreczy & Rácz var. sinaloensis (Urb.) Staszk. var. latifolius (Florin) Staszk. 818
Debreczy & Rácz 744 Podocarpus angustifolius Griseb. var. leonii (Cara-
Pinus yunnanensis Franch. 609, 614, 662, 760, bia) Staszk. 807
782, 783 Podocarpus angustifolius Griseb. var. leonii
Pinus yunnanensis Franch. var. pygmaea (J. R. Xue) (Carabia) Staszk. f. victorinianus (Carabia)
J. R. Xue 783 Staszk. 807
Pinus yunnanensis Franch. var. tenuifolia W. C. Podocarpus angustifolius Parl. 876
Cheng & Y. W. Law 782, 783 Podocarpus annamiensis N. E. Gray 867, 869
Pinus yunnanensis Franch. var. yunnanensis Podocarpus annamiensis N. E. Gray var. hainanensis
782, 783 Gaussen 869
Platycladus Spach 36, 46, 48, 514, 793, 794 Podocarpus aracensis de Laub. & Silba 803, 809
Platycladus chengii (Bordères & Gaussen) A. V. Bo- Podocarpus archboldii N. E. Gray 800, 810, 826
brov 793
Podocarpus archboldii N. E. Gray var. crassiramosus Podocarpus costalis C. Presl var. taiwanensis Gaus-
N. E. Gray 883 sen 824
Podocarpus argotaenia Hance 168, 169 Podocarpus costaricensis de Laub. ex Silba 802,
Podocarpus aristulatus Parl. 807, 808, 818, 832 824, 825
Podocarpus aristulatus Parl. var. buchii (Urb.) Podocarpus crassigemma de Laub. 800, 825, 826
Silba 818 Podocarpus cumingii Parl. 320
Podocarpus aspleniifolius Labill. 544 Podocarpus cunninghamii Colenso 322, 380, 505,
Podocarpus atjehensis (Wasscher) de Laub. ex 507, 550, 787, 801, 806, 826, 827, 908
Silba 797, 799, 811 Podocarpus cupressinus R. Br. ex G. Benn. var. cur­
Podocarpus ballivianensis Silba 873 vulus Miq. 325
Podocarpus barretoi de Laub. & Silba 813 Podocarpus dacrydiifolius Wasscher 318 1103
Podocarpus beecherae de Laub. 870, 871 Podocarpus dacrydioides A. Rich. 317, 321
Podocarpus biformis Hook. 378 Podocarpus dawei Stapf 137
Podocarpus borneensis de Laub. 798, 799, 812 Podocarpus decipiens N. E. Gray 869
Podocarpus bracteatus Blume 798, 812, 813, 883 Podocarpus decumbens N. E. Gray 801, 828
Podocarpus brasiliensis de Laub. 804, 813, 814 Podocarpus deflexus Ridl. 798, 828, 829
Podocarpus brassii Pilg. 787, 799, 814, 815, 816 Podocarpus degeneri (N. E. Gray) de Laub. ex
Podocarpus brassii Pilg. var. brassii 787, 815, 816, Silba 355, 869
1069 Podocarpus dispermus C. T. White 787, 800, 829,
Podocarpus brassii Pilg. var. humilis de Laub. 815, 830, 953
816, 817 Podocarpus distichus J. T. Buchholz 917
Podocarpus brevifolius (Stapf) Foxw. 326, 349, Podocarpus distichus J. T. Buchholz var. maialis
787, 799, 817, 818, 1032 J. T. Buchholz 917
Podocarpus brownii C. E. Bertrand 830 Podocarpus drouynianus F. Muell. 800, 830, 831,
Podocarpus buchholzii de Laub. 809, 887, 888 864
Podocarpus buchholzii de Laub. var. neblinensis Podocarpus ekmanii Urb. 802, 831, 832
Silba 887 Podocarpus elatus R. Br. ex Endl. 232, 788, 800,
Podocarpus buchii Urb. 802, 818, 832 804, 807, 832, 833, 834
Podocarpus buchii Urb. var. latifolius Florin 818, Podocarpus elongatus (Aiton) L’Herit. ex
819 Pers. 795, 796, 804, 834, 835
Podocarpus capuronii de Laub. 796, 819, 820 Podocarpus ensiculus Melville 843
Podocarpus capuronii de Laub. var. woltzii (Gaus- Podocarpus epiphyticus de Laub. & Silba 868, 869
sen) Silba 819 Podocarpus expansus (de Laub.) Whitmore 322
Podocarpus cardenasii J. T. Buchholz & N. E. Gray Podocarpus falcatus (Thunb.) Endl. 138
839 Podocarpus falciformis Parl. 363, 364
Podocarpus celatus de Laub. 803, 820, 821 Podocarpus fasciculus de Laub. 797, 805, 835, 836,
Podocarpus celebicus Warb. 880 903
Podocarpus chinensis Wall. ex J. Forbes 859 Podocarpus ferrugineus G. Benn. ex D. Don 916
Podocarpus chinensis Wall. ex J. Forbes var. wardii Podocarpus ferruginoides R. H. Compton 917
de Laub. & Silba 859 Podocarpus filicifolius N. E. Gray 942
Podocarpus chingianus (N. E. Gray) S. Y. Hu 787, Podocarpus fleuryi Hickel 518
797, 821, 822, 858 Podocarpus formosensis Dummer 521
Podocarpus cinctus Pilg. 318 Podocarpus formosensis Dummer var. koshuensis
Podocarpus colliculatus (N. E. Gray) de Laub. 904 (Kaneh.) Merr. & Yamam. 522
Podocarpus compactus Wasscher 319 Podocarpus gaussenii Woltz 138, 139
Podocarpus comptonii J. T. Buchholz 937 Podocarpus gibbsiae N. E. Gray 349, 799, 837
Podocarpus confertus de Laub. 799, 822 Podocarpus glaucus Foxw. 548, 797, 799, 837, 838,
Podocarpus coriaceus Rich. 802, 823, 910 847
Podocarpus costalis C. Presl 787, 797, 804, 824, Podocarpus globulus de Laub. 798, 838
886 Podocarpus glomeratus D. Don 803, 839, 840
Podocarpus gnidioides Carrière 801, 840, 841
 Podocarpus gnidioides Carrière var. caespitosus Podocarpus macrocarpus de Laub. 798, 856, 857
Pancher ex Carrière 840 Podocarpus macrophyllus (Thunb.) Sweet 169,
Podocarpus gracilimus Stapf 138, 139 266, 795, 797, 805, 821, 836, 837, 857, 858, 866,
Podocarpus gracilior Pilg. 139 867, 1005, 1070
Podocarpus grayae de Laub. 788, 800, 841 Podocarpus macrophyllus (Thunb.) Sweet var.
Podocarpus guatemalensis Standl. 802, 804, 842, chingii N. E. Gray 821, 858
843 Podocarpus macrophyllus (Thunb.) Sweet f. grandi­
Podocarpus guatemalensis Standl. var. allenii folius Pilg. 835, 836
(Standl.) J. T. Buchholz & N. E. Gray 842 Podocarpus macrophyllus (Thunb.) Sweet var.
Podocarpus guatemalensis Standl. var. pinetorum liukiuensis Warb. 835, 836
1104 (Bartlett) J. T. Buchholz & N. E. Gray 842 Podocarpus macrophyllus (Thunb.) Sweet var. mac­
Podocarpus hallii Kirk 826, 827 rophyllus 789, 858, 866
Podocarpus harmsianus Pilg. 918 Podocarpus macrophyllus (Thunb.) Sweet subsp.
Podocarpus henkelii Stapf ex Dallim. & A. B. Jacks. maki (Siebold & Zucc.) Pilg. 859
136, 788, 796, 843, 844 Podocarpus macrophyllus (Thunb.) Sweet var. maki
Podocarpus hispaniolensis de Laub. 802, 844 Siebold & Zucc. 858, 859, 866
Podocarpus humbertii de Laub. 796, 845, 846 Podocarpus macrophyllus (Thunb.) Sweet var. nakaii
Podocarpus idenburgensis N. E. Gray 852 (Hayata) H. L. Li & Keng 866
Podocarpus imbricatus Blume 317, 324 Podocarpus macrophyllus (Thunb.) Sweet var. pilira­
Podocarpus imbricatus Blume var. kinabaluensis mulus Z. X. Chen & Z. Q. Li 858
Wasscher 325 Podocarpus macrostachyus Parl. 873, 874
Podocarpus indonesiensis de Laub. & Silba 889, Podocarpus madagascariensis Baker 796, 859,
890 860
Podocarpus ingensis de Laub. 873, 874 Podocarpus madagascariensis Baker var. madagas­
Podocarpus insularis de Laub. 846, 847 cariensis 860
Podocarpus kawaii Hayata 324 Podocarpus madagascariensis Baker var. procerus
Podocarpus koordersii Pilg. ex Koord. & Vale- de Laub. 860
ton 890 Podocarpus madagascariensis Baker var. rotundus
Podocarpus koshunensis (Kaneh.) Kaneh. 522 L. Laurent 861
Podocarpus ladei F. M. Bailey 919 Podocarpus magnifolius J. T. Buchholz &
Podocarpus lambertii Klotzsch ex Endl. 190, 803, N. E. Gray 802, 803, 820, 821, 861, 862
847, 848, 849, 896, 908, 909 Podocarpus maki (Siebold & Zucc.) Gaussen 859
Podocarpus lambertii Klotzsch ex Endl. var. tran­ Podocarpus mannii Hook. f. 141
siens Pilg. 908 Podocarpus matudae Lundell 789, 802, 804, 825,
Podocarpus latifolius (Thunb.) R. Br. ex 862, 863, 981
Mirb. 139, 523, 788, 796, 805, 849, 850 Podocarpus matudae Lundell var. jaliscanus de
Podocarpus latifolius (Thunb.) R. Br. ex Mirb. var. Laub. & Silba 862, 863
latior Pilg. 849 Podocarpus matudae Lundell var. macrocarpus
Podocarpus latifolius Wall. 523 J. T. Buchholz & N. E. Gray 862, 863
Podocarpus latior (Pilg.) Gaussen 849 Podocarpus matudae Lundell var. reichei (J. T. Buch-
Podocarpus laubenfelsii Tiong 350, 788, 797, 798, holz & N. E. Gray) de Laub. & Silba 862
850, 851 Podocarpus micropedunculatus de Laub. 799,
Podocarpus lawrencei Hook. f. 362, 516, 545, 789, 863, 864
800, 804, 840, 851, 852 Podocarpus milanjianus Rendle 142, 439, 789,
Podocarpus ledermannii Pilg. 800, 852, 853 796, 844, 864, 865, 1024
Podocarpus leonii Carabia 807 Podocarpus minus (Carrière) Parl. 938
Podocarpus leptophyllus Wasscher 351, 352 Podocarpus montanus (Humb. & Bonpl. ex Willd.)
Podocarpus levis de Laub. 798, 799, 800, 853, 854 Lodd. ex Britton 920
Podocarpus longifoliolatus Pilg. 801, 854 Podocarpus montanus (Willd.) Lodd. var. densi­
Podocarpus lophatus de Laub. 797, 855 folius J. T. Buchholz & N. E. Gray 920
Podocarpus lucienii de Laub. 801, 855, 856
Podocarpus montanus (Willd.) Lodd. var. diversi­ Podocarpus oleifolius D. Don var. macrostachyus
folius Dallim. & A. B. Jacks. 920 (Parl.) J. T. Buchholz & N. E. Gray 873
Podocarpus montanus (Willd.) Lodd. var. meriden­ Podocarpus oleifolius D. Don var. trujillensis
sis J. T. Buchholz & N. E. Gray 920 J. T. Buchholz & N. E. Gray 873
Podocarpus monteverdeensis de Laub. 873, 874 Podocarpus palawanensis de Laub. & Silba 798,
Podocarpus nagi (Thunb.) Pilg. var. koshunensis 874
Kaneh. 522 Podocarpus pallidus N. E. Gray 801, 875, 876
Podocarpus nakaii Hayata 172, 473, 789, 796, 866 Podocarpus palustris J. T. Buchholz 938
Podocarpus nankoensis Hayata 521 Podocarpus papuanus Ridl. 319, 325
Podocarpus neriifolius D. Don 169, 173, 176, 251, Podocarpus parlatorei Pilg. 803, 876, 892
270, 324, 797, 798, 799, 800, 801, 805, 811, 813, Podocarpus pectinatus Pancher ex Brongn. & 1105
842, 847, 853, 858, 867, 868, 869, 886, 891, 903, Gris 129
904, 943, 984 Podocarpus pendulifolius J. T. Buchholz &
Podocarpus neriifolius D. Don var. atjehensis Was­ N. E. Gray 804, 877, 878, 879
scher 811 Podocarpus perrieri Gaussen & Woltz 796, 879,
Podocarpus neriifolius D. Don var. bracteatus 880
(Blume) Wasscher 812 Podocarpus philippinensis Foxw. 890
Podocarpus neriifolius D. Don var. brevifolius Podocarpus pilgeri Foxw. 169, 173, 270, 797, 798,
Stapf 817 799, 807, 815, 855, 880, 881, 886, 975
Podocarpus neriifolius D. Don var. decipiens Podocarpus pilgeri Foxw. var. thailandensis Gaus-
(N. E. Gray) Silba 869 sen 880
Podocarpus neriifolius D. Don var. degeneri Podocarpus pinetorum Bartlett 842
N. E. Gray 867, 868, 869 Podocarpus pittieri J. T. Buchholz & N. E. Gray 892
Podocarpus neriifolius D. Don var. neriifolius Podocarpus polyanthus (Wasscher) Gaussen 868
789, 868 Podocarpus polyspermus de Laub. 801, 881, 882
Podocarpus neriifolius D. Don var. penibukanensis Podocarpus polystachyus R. Br. ex Endl. 790, 797,
Silba 869 798, 799, 800, 805, 824, 875, 882, 883
Podocarpus neriifolius D. Don var. polyanthus Was­ Podocarpus polystachyus R. Br. ex Endl. var. rigidus
scher 868 Wasscher 812
Podocarpus neriifolius D. Don var. ridleyi Was­ Podocarpus polystachyus R. Br. ex Endl. var.
scher 886 ­thevetiifolius (Zipp. ex Blume) Silba 882
Podocarpus neriifolius D. Don var. staintonii Podocarpus pseudobracteatus de Laub. 800, 883,
Silba 869 884
Podocarpus neriifolius D. Don var. teysmannii Podocarpus purdieanus Hook. 802, 884, 885
(Miq.) Wasscher 906 Podocarpus ramosii R. R. Mill 798, 885, 886
Podocarpus neriifolius D. Don var. timorensis Was­ Podocarpus reichei J. T. Buchholz & N. E. Gray
scher 889 862, 863
Podocarpus nivalis Hook. 378, 379, 502, 790, 801, Podocarpus ridleyi (Wasscher) N. E. Gray 798,
804, 840, 869, 870 886, 887, 898
Podocarpus novae-caledoniae Vieill. 242, 526, Podocarpus roraimae Pilg. 803, 887, 888, 901, 906
790, 801, 870, 871, 904 Podocarpus rospigliosii Pilg. 941
Podocarpus novae-caledoniae Vieill. var. colliculatus Podocarpus rostratus L. Laurent 796, 819, 880,
N. E. Gray 904 888, 889
Podocarpus nubigenus Lindl. 370, 607, 790, 803, Podocarpus rostratus L. Laurent var. perrieri (Gaus-
804, 872, 873, 944 sen & Woltz) Silba 879
Podocarpus oleifolius D. Don 802, 804, 821, 843, Podocarpus rotundus de Laub. 885, 886
873, 874 Podocarpus rubens de Laub. 889
Podocarpus oleifolius D. Don var. costaricensis Podocarpus rubens de Laub. var. pabinamaensis
J. T. Buchholz & N. E. Gray 873 Silba 889
Podocarpus oleifolius D. Don var. equadorensis Podocarpus rubens de Laub. var. sumatrana
Silba 873 Silba 889
 Podocarpus rumphii Blume 790, 797, 798, 799, Podocarpus transiens (Pilg.) de Laub. ex Silba var.
857, 890, 891 harleyi Silba 908
Podocarpus rumphii Blume var. arbainii Silba 890 Podocarpus trinitensis J. T. Buchholz & N. E. Gray
Podocarpus rumphii Blume var. aruensis Silba 890 802, 909, 910
Podocarpus rusbyi J. T. Buchholz & N. E. Gray Podocarpus urbanii Pilg. 802, 910, 911, 912
803, 892 Podocarpus usambarensis Pilg. 141
Podocarpus salicifolius Klotzsch & H. Karst. ex Podocarpus ustus (Vieill.) Brongn. & Gris 539
Endl. 803, 878, 892, 893 Podocarpus utilior Pilg. 918
Podocarpus salignus D. Don 370, 791, 803, 804, Podocarpus victorinianus Carabia 807, 808
893, 894 Podocarpus vieillardii Parl. 327
1106 Podocarpus salomoniensis Wasscher 800, 894 Podocarpus vitiensis Seem. 937, 942
Podocarpus sellowii Klotzsch ex Endl. 190, 803, Podocarpus wallichianus C. Presl 523
849, 895, 896 Podocarpus woltzii Gaussen 819
Podocarpus sellowii Klotzsch ex Endl. var. angusti­ Prumnopitys Phil. 18, 41, 51, 52, 913, 918, 922, 923,
folius Pilg. 896, 897 942, 953
Podocarpus sellowii Klotzsch ex Endl. var. sel­ Prumnopitys amara (Blume) de Laub. 952
lowii 896 Prumnopitys andina (Poepp. ex Endl.) de Laub.
Podocarpus smithii de Laub. 800, 897, 898, 919 370, 792, 913, 914, 915, 922
Podocarpus spathoides de Laub. 798, 799, 800, Prumnopitys andina (Poepp. ex Endl.) de Laub.
898 subsp. blijdensteinii Silba 914
Podocarpus spathoides de Laub. var. solomonensis Prumnopitys elegans Phil. 913, 914
Silba 898 Prumnopitys exigua de Laub. 913, 915, 922
Podocarpus spicatus Poepp. 914 Prumnopitys ferruginea (G. Benn. ex D. Don) de
Podocarpus spinulosus (Sm.) R. Br. ex Mirb. 791, Laub. 147, 322, 346, 378, 380, 507, 549, 827,
800, 899 908, 914, 916, 917
Podocarpus sprucei Parl. 791, 803, 900 Prumnopitys ferruginoides (R. H. Compton) de
Podocarpus standleyi J. T. Buchholz & N. E. Gray  921 Laub. 202, 205, 352, 504, 792, 914, 917
Podocarpus steupii Wasscher 326 Prumnopitys harmsiana (Pilg.) de Laub. 913, 918,
Podocarpus steyermarkii J. T. Buchholz & N. E. 919
Gray 803, 888, 901, 902, 906 Prumnopitys ladei (F. M. Bailey) de Laub. 792,
Podocarpus subtropicalis de Laub. 797, 805, 867, 914, 919, 920
868, 903 Prumnopitys montana (Humb. & Bonpl. ex Willd.)
Podocarpus subtropicalis de Laub. var. medogensis de Laub. 913, 915, 920, 921, 922
Silba 903 Prumnopitys spicata (Poepp.) Molloy & Muñoz-
Podocarpus sylvestris J. T. Buchholz 156, 214, 904 Schick 914
Podocarpus tepuiensis J. T. Buchholz & N. E. Gray Prumnopitys standleyi (J. T. Buchholz & N. E. Gray)
803, 888, 901, 905, 906 de Laub. 913, 921, 922
Podocarpus teysmannii Miq. 797, 798, 906 Prumnopitys taxifolia (Banks & Sol. ex D. Don) de
Podocarpus thevetiifolius Zipp. ex Blume 882 Laub. 147, 322, 346, 378, 507, 549, 913, 916, 917,
Podocarpus tixieri Gaussen 880 922, 923
Podocarpus tixieri Gaussen ex Silba 880 Pseudolarix Gordon 15, 17, 39, 49, 50, 924
Podocarpus totara G. Benn. ex D. Don 18, 26, 139, Pseudolarix amabilis (J. Nelson) Rehd. 273, 792,
147, 322, 346, 507, 549, 791, 801, 804, 827, 907, 924, 925
908, 916, 923, 1070 Pseudolarix kaempferi (Lindl.) Gordon 924
Podocarpus totara G. Benn. ex D. Don var. hallii Pseudolarix pourtetii Ferré 924
(Kirk) Pilg. 826 Pseudotaxus W. C. Cheng 41, 53, 927
Podocarpus totara G. Benn. ex D. Don var. waiho­ Pseudotaxus chienii (W. C. Cheng) W. C. Cheng
ensis Wardle 907 289, 792, 927, 928
Podocarpus transiens (Pilg.) de Laub. ex Silba Pseudotaxus chienii (W. C. Cheng) W. C. Cheng
803, 908 subsp. liana (Silba) Silba 927
Pseudotaxus liana Silba 927, 928
Pseudotsuga Carrière 17, 38, 49, 50, 267, 633, 687, Retinispora formosensis (Matsum.) A. V. Bobrov &
761, 929, 934 Melikyan 274
Pseudotsuga argyrophylla (Chun & Kuang) Gre- Retinispora lawsoniana (A. Murray bis) A. V. Bo-
guss 252 brov & Melikyan 276
Pseudotsuga brevifolia W. C. Cheng & L. K. Fu 934 Retinispora obtusa Siebold & Zucc. 274, 278
Pseudotsuga davidiana Bertrand 474 Retinispora pisifera Siebold & Zucc. 280
Pseudotsuga flahaultii Flous 933 Retinispora taiwanensis (Masam. & S. Suzuki)
Pseudotsuga forrestii Craib 934, 935 A. V. Bobrov & Melikyan 280
Pseudotsuga gausseni Flous 935 Retrophyllum C. N. Page 39, 51, 52, 937
Pseudotsuga japonica (Shiras.) Beissn. 83, 280, Retrophyllum comptonii (J. T. Buchholz)
522, 929, 930, 961, 1019 C. N. Page 161, 937, 938, 962 1107
Pseudotsuga macrocarpa (Vasey) Mayr 247, 929, Retrophyllum minus (Carrière) C. N. Page 18,
931 340, 351, 937, 938, 939, 940, 962
Pseudotsuga menziesii (Mirb.) Franco 25, 49, 61, Retrophyllum piresii (Silba) C. N. Page 937, 940
66, 88, 99, 113, 247, 277, 292, 295, 304, 312, 493, Retrophyllum rospigliosii (Pilg.) C. N. Page 937,
564, 568, 600, 670, 676, 680, 698, 705, 715, 749, 940, 941
929, 932, 949, 951, 961, 973, 993, 999, 1016 Retrophyllum vitiense (Seem.) C. N. Page 810,
Pseudotsuga menziesii (Mirb.) Franco var. flahaultii 937, 942, 943
(Flous) Silba 933
Pseudotsuga menziesii (Mirb.) Franco subsp. glauca
S
(Beissn.) E. Murray 933
Pseudotsuga menziesii (Mirb.) Franco var. glauca Sabina Mill. 381
(Beissn.) Franco 73, 78, 116, 125, 308, 566, 593, Sabina ashei (J. T. Buchholz) A. V. Bobrov & Me-
932, 933, 993 likyan 387
Pseudo­tsuga menziesii (Mirb.) Franco subsp. macro­ Sabina convallium (Rehd. & E. H. Wilson) W. C.
carpa (Vasey) E. Murray 931 Cheng & L. K. Fu var. microsperma W. C.
Pseudotsuga menziesii (Mirb.) Franco var. men­ Cheng & L. K. Fu 404
ziesii 933, 961 Sabina microsperma (W. C. Cheng & L. K. Fu)
Pseudotsuga menziesii (Mirb.) Franco var. oaxacana W. C. Cheng & L. K. Fu 404
Debreczy & Rácz 933 Sabina pingii (W. C. Cheng) W. C. Cheng & W. T.
Pseudotsuga sinensis Dode 95, 176, 248, 251, 270, Wang var. wilsonii (Rehd.) W. C. Cheng &
287, 476, 519, 522, 587, 858, 929, 933, 934, 935, L. K. Fu 438
955, 975, 1015, 1032 Sabina silicicola Small 470
Pseudotsuga sinensis Dode var. brevifolia Sabina vulgaris Antoine 381
(W. C. Cheng & L. K. Fu) Farjon & Silba 935 Sabina vulgaris Antoine var. erectopatens
Pseudotsuga sinensis Dode var. forrestii (Craib) W. C. Cheng & L. K. Fu 396
Silba 935 Sabina vulgaris Antoine var. yulinensis T. C. Chang
Pseudotsuga sinensis Dode var. gaussenii (Flous) & C. G. Chen 448
Silba 929, 935 Sabinella Nakai 381
Pseudotsuga sinensis Dode var. sinensis 929, 934, Sabinella phoenicea (L.) Nakai 381
961 Salisburyodendron A. V. Bobrov & Melikyan 143
Pseudotsuga sinensis Dode var. wil­soniana (Hayata) Salisburyodendron australis (D. Don) A. V. Bobrov
L. K. Fu & Nan Li & Melikyan 143, 146
Pseudotsuga wilsoniana Hayata 934 Salisburyodendron corbassonii (de Laub.) A. V. Bo-
Pseudotsuga xichangensis C. T. Kuan & L. J. Zhou   935 brov & Melikyan 161
Pseudotsuga shaanxiensis S. Z. Qu & K. Y. Wang   935 Salisburyodendron lanceolata (Warb.) A. V. Bobrov
& Melikyan 155
Salisburyodendron montana (de Laub.) A. V. Bobrov
R
& Melikyan 160
Retinispora Siebold & Zucc. 274 Salisburyodendron moorei (Lindl.) A. V. Bobrov &
Melikyan 161
 Salisburyodendron ovata (Vieill.) A. V. Bobrov & Tassilicyparis dupreziana (A. Camus) A. V. Bobrov
Melikyan 163 & Melikyan 290, 301
Salisburyodendron ovata (Vieill.) A. V. Bobrov Taxaceae Gray 16, 19, 20, 22, 24, 33, 41, 43, 45, 52,
& Melikyan subsp. hypoleuca (C. Moore ex 168, 222, 223, 251, 539, 927, 969, 998, 1054, 1056,
Henkel & W. Hochst.) A. V. Bobrov & Me- 1059, 1060, 1061, 1062
likyan 163 Taxodium Rich. 15, 18, 35, 46, 47, 376, 922, 956
Saxegothaea Lindl. 39, 51, 52, 944 Taxodium ascendens Brongn. 958
Saxegothaea conspicua Lindl. 370, 872, 944, 945, Taxodium distichum (L.) Rich. 282, 313, 513, 671,
962 679, 758, 948, 956, 957, 958, 959, 964
Schubertia Mirb. 956 Taxodium distichum (L.) Rich. var. ascendens
1108 Schubertia disticha (L.) Mirb. 956 (Brongn.) Mast. 958
Sciadopityaceae Luerss. 19, 22, 41, 43, 52, 946, Taxodium distichum (L.) Rich. var. distichum 958
1034 Taxodium distichum (L.) Rich. var. imbricarium
Sciadopitys Siebold & Zucc. 16, 41, 46, 52, 316, (Nutt.) Sarg. 375, 957, 958
391, 423, 437, 448, 457, 946, 947 Taxodium distichum (L.) Rich. var. mucronatum
Sciadopitys verticillata (Thunb.) Siebold & (Ten.) A. Henry 958
Zucc. 19, 83, 280, 946, 962, 996, 1005, 1019 Taxodium distichum (L.) Rich. subsp. nutans
Sequoia Endl. 18, 22, 34, 46, 47, 232, 948 (­Aiton) E. Murray 958
Sequoia sempervirens (D. Don) Endl. 19, 46, 277, Taxodium mucronatum Ten. 956, 958, 959, 964
948, 949, 963, 973, 993, 999, 1037 Taxodium sempervirens D. Don 948
Sequoiadendron J. T. Buchholz 22, 34, 46, 47, 232, Taxus L. 19, 20, 26, 42, 52, 53, 495, 539, 545, 944,
275, 950 946, 969, 977, 980, 981, 983, 984, 1005
Sequoiadendron giganteum (Lindl.) J. T. Buch- Taxus baccata L. 72, 107, 109, 255, 592, 602, 734,
holz 19, 23, 247, 690, 696, 950, 951, 963 964, 965, 969, 970, 971, 972, 975, 978, 984, 985,
Squamataxus J. Nelson 944 1054
Squamataxus albertiana J. Nelson 944 Taxus baccata L. subsp. brevifolia (Nutt.) Pilg. 972
Stachycarpus Tiegh. 913 Taxus baccata L. subsp. canadensis (Marshall)
Stachycarpus amarus (Blume) Gaussen 952 Pilg. 973
Stachycarpus andinus (Poepp. ex Endl.) Tiegh. 913 Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Stegocedrus Doweld 503 Pilg. 978
Stegocedrus austrocaledonica (Brongn. & Gris) Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Doweld 503, 504 Pilg. var. chinensis Pilg. 974
Stegocedrus chevalieri (J. T. Buchholz) Doweld 506 Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Stegocedrus yateensis (Guillaumin) Doweld 508 Pilg. var. latifolia Pilg. 978
Steinhauera Presl 950 Taxus baccata L. subsp. floridana (Nutt. ex Chapm.)
Steinhauera gigantea (Lindl.) Kuntze ex Voss 950 Pilg. 979
Strobus Opiz 608 Taxus baccata L. var. microcarpa Trautv. 978
Strobus weymouthiana Opiz 608 Taxus baccata L. var. minor Michx. 973
Sundacarpus (J. T. Buchholz & N. E. Gray) Taxus baccata L. subsp. wallichiana (Zucc.) Pilg.
C. N. Page 41, 51, 52, 366, 952 983, 984
Sundacarpus amarus (Blume) C. N. Page 148, 321, Taxus biternata Spjut 978
343, 350, 365, 366, 524, 851, 919, 952, 953, 963 Taxus brevifolia Nutt. 277, 715, 951, 965, 970, 972,
973, 993
Taxus brevifolia Nutt. var. polychaeta Spjut 972
T
Taxus brevifolia Nutt. var. reptaneta Spjut 972
Taiwania Hayata 34, 46, 47, 954, 955 Taxus caespitosa Nakai 978
Taiwania cryptomerioides Hayata 248, 275, 280, Taxus caespitosa Nakai var. angustifolia Spjut 979
287, 954, 955, 963, 964 Taxus caespitosa Nakai var. latifolia (Pilg.)
Taiwania flousiana Gaussen 954 ­Spjut 978
Taiwania yunnanensis Koidz. 954 Taxus canadensis Marshall 62, 970, 972, 973, 974
Tassilicyparis A. V. Bobrov & Melikyan 290
Taxus canadensis Marshall var. floridana (Nutt. ex Taxus nucifera L. 998, 1004
Chapm.) Silba 979 Taxus obscura Spjut 983
Taxus canadensis Marshall var. minor (Michx.) Taxus phytonii Spjut 983
­Spjut 973 Taxus speciosa Florin 981, 982
Taxus celebica (Warb.) H. L. Li 983, 984 Taxus spinulosa Sm. 899
Taxus chienii W. C. Cheng 927 Taxus suffnessii Spjut 983
Taxus chinensis (Pilg.) Rehd. 64, 82, 169, 176, 263, Taxus sumatrana (Miq.) de Laub. 983, 984
270, 289, 476, 522, 858, 970, 974, 975, 982, 984, Taxus umbraculifera (Siebold ex Endl.) Lawson var.
1008, 1032 microcarpa (Trautv.) Spjut 978
Taxus chinensis (Pilg.) Rehd. var. mairei (Lemée & Taxus umbraculifera (Siebold ex Endl.) Lawson var.
Lév.) W. C. Cheng & L. K. Fu 981 nana (Rehd.) Spjut 979 1109
Taxus chinensis (Pilg.) Rehd. var. yunnanensis Taxus verticillata Thunb. 946
(W. C. Cheng & L. K. Fu) L. K. Fu 983 Taxus wallichiana Zucc. 270, 563, 955, 970, 975,
Taxus communis J. Nelson 969 976, 977, 982, 983, 984, 985, 1059, 1061
Taxus contorta Griff. 970, 976, 977, 984, 985 Taxus wallichiana Zucc. var. chinensis (Pilg.)
Taxus contorta Griff. var. mucronata Spjut 983 ­Florin 974
Taxus cuspidata Siebold & Zucc. 96, 573, 970, Taxus wallichiana Zucc. var. mairei (Lemée & Lév.)
975, 977, 978, 1005, 1071 L. K. Fu & Nan Li 981
Taxus cuspidata Siebold & Zucc. var. caespitosa Taxus wallichiana Zucc. var. yunnanensis
(Nakai) Q. L. Wang 978 (W. C. Cheng & L. K. Fu) C. T. Kuan 983
Taxus cuspidata Siebold & Zucc. var. chinensis Taxus yunnanensis W. C. Cheng & L. K. Fu 976,
(Pilg.) C. K. Schneid. 974 983, 1002, 1014
Taxus cuspidata Siebold & Zucc. var. cuspida­ Tetraclinis Mast. 36, 46, 47, 514, 986, 988
ta 965, 978 Tetraclinis articulata (Vahl) Mast. 26, 282, 301,
Taxus cuspidata Siebold & Zucc. var. latifolia (Pilg.) 965, 986, 987, 1060
Nakai 978 Thalamia Spreng. 544
Taxus cuspidata Siebold & Zucc. var. microcarpa Thalamia aspleniifolia (Labill.) Spreng. 544
(Trautv.) Kolesn. 978 Thuja L. 35, 46, 48, 220, 514, 793, 932, 989
Taxus cuspidata Siebold & Zucc. var. nana hort. ex Thuja articulata Vahl 986
Rehd. 979 Thuja chengii 793
Taxus elongata Aiton 795, 834 Thuja chilensis D. Don 220
Taxus falcata Thunb. 136, 138 Thuja cupressoides L. 1020, 1022
Taxus floridana Nutt. ex Chapm. 970, 979, 980, Thuja dolabrata Thunb. ex L. f. 996, 997
1006 Thuja koraiensis Nakai 965, 966, 989, 990
Taxus florinii Spjut 983 Thuja lineata Poir. 375
Taxus fuana Nan Li & R. R. Mill 976, 984 Thuja occidentalis L. 779, 989, 991, 993
Taxus globosa Schltdl. 583, 970, 980, 981 Thuja orientalis L. 793, 1057
Taxus globosa Schltdl. var. floridana (Nutt. ex Thuja pensilis Staunton ex D. Don 375
Chapm.) Spjut 979 Thuja plicata Donn ex D. Don 61, 88, 113, 247,
Taxus harringtonii Knight ex J. Forbes 261, 265, 493, 600, 698, 715, 966, 973, 989, 992, 993, 1029
266 Thuja sphaeroidea Spreng. 274
Taxus kingstonii Spjut 981 Thuja standishii (Gordon) Carrière 94, 123, 989,
Taxus latifolia Thunb. 849 993, 994, 996, 1013
Taxus macrophylla Thunb. 795, 858 Thuja sutchuenensis Franch. 966, 989, 994, 1063
Taxus mairei (Lemée & Lév.) S. Y. Hu ex Thujopsis Siebold & Zucc. ex Endl. 35, 46, 47, 996
T. S. Liu 970, 981, 982, 984 Thujopsis dolabrata (Thunb. ex L. f.) Siebold &
Taxus mairei (Lemée & Lév.) S. Y. Hu ex T. S. Liu Zucc. 966, 994, 996
var. speciosa (Florin) Spjut 981 Thujopsis dolabrata (Thunb. ex L. f.) Siebold &
Taxus ×media Rehd. 978 Zucc. var. dolabrata 966, 997, 1071
Taxus minor (Michx.) Britton ex Small & Vail 973 Thujopsis dolabrata (Thunb. ex L. f.) Siebold &
Taxus montana Humb. & Bonpl. ex Willd. 920 Zucc. var. hondae Makino 996, 997, 1013
 Thujopsis standishii Gordon 993 Tsuga chinensis (Franch.) E. Pritz. var. formosana
Titanodendron A. V. Bobrov & Melikyan 185 (Hayata) H. L. Li & H. Keng 1011
Titanodendron hunsteinii (K. Schum.) A. V. Bobrov Tsuga chinensis (Franch.) E. Pritz. var. forrestii
& Melikyan 185, 202 (Downie) Silba 1014
Titanodendron klinkii (Lauterb.) A. V. Bobrov & Tsuga chinensis (Franch.) E. Pritz. var. oblongi­
Melikyan 202 squamata W. C. Cheng & L. K. Fu 1012
Titanodendron schumanniana (Warb.) A. V. Bobrov Tsuga chinensis (Franch.) E. Pritz. subsp. patens
& Melikyan 202 (Downie) E. Murray 1011
Torreya Arn. 20, 42, 53, 998, 1006 Tsuga chinensis (Franch.) E. Pritz. var. patens
Torreya ascendens Nakai ex Uyeki 1004 (Downie) L. K. Fu & Nan Li 1011
1110 Torreya californica Torr. 966, 998, 999 Tsuga chinensis (Franch.) E. Pritz. var. robusta
Torreya fargesii Franch. 998, 999, 1000 W. C. Cheng & L. K. Fu 1012
Torreya fargesii Franch. var. fargesii 1000, 1001 Tsuga chinensis (Franch.) E. Pritz. var. tchekiangen­
Torreya fargesii Franch. var. yunnanensis (W. C. sis (Flous) W. C. Cheng & L. K. Fu 1011
Cheng & L. K. Fu) N. Kang 1000, 1001 Tsuga chinensis (Franch.) E. Pritz. subsp. wardii
Torreya grandis Fortune ex Lindl. 289, 473, 955, (Downie) E. Murray 1013
998, 1000, 1001, 1002 Tsuga diversifolia (Maxim.) Mast. 94, 100, 123,
Torreya grandis Fortune ex Lindl. var. fargesii 489, 557, 574, 994, 996, 1007, 1008, 1012, 1013
(Franch.) Silba 1000 Tsuga diversifolia (Maxim.) Mast. subsp. blaring­
Torreya grandis Fortune ex Lindl. var. gran­ hemii (Flous) E. Murray 1012
dis 1000, 1002 Tsuga douglasii (Sabine ex D. Don) Lindl. var. glau­
Torreya grandis Fortune ex Lindl. var. jiulong­shan­ ca Beissn. 933
ensis Z. Y. Li, Z. C. Tang & N. Kang 1000, 1003 Tsuga dumosa (D. Don) Eichler 75, 77, 84, 110,
Torreya grandis Fortune ex Lindl. var. yunnanensis 121, 297, 487, 495, 563, 569, 601, 602, 780, 934,
(W. C. Cheng & L. K. Fu) Silba 1001 955, 1007, 1008, 1013, 1014, 1015
Torreya jackii Chun 998, 1003, 1004 Tsuga dumosa (D. Don) Eichler subsp. leptophylla
Torreya nucifera (L.) Siebold & Zucc. 83, 285, 930, (Hand.-Mazz.) E. Murray 1013
967, 998, 1004, 1005 Tsuga dumosa (D. Don) Eichler var. yunnanensis
Torreya nucifera (L.) Siebold & Zucc. var. grandis (Franch.) Silba 1013
(Fortune ex Lindl.) Pilg. 1002 Tsuga dura Downie 1013
Torreya parvifolia T. P. Yi, L. Yang & T. L. Long 1002 Tsuga formosana Hayata 1011
Torreya taxifolia Arn. 980, 998, 1005, 1006, 1053, Tsuga forrestii Downie 122, 967, 1007, 1008, 1014,
1061 1015
Torreya yunnanensis W. C. Cheng & L. K. Fu 1001 Tsuga heterophylla (Raf.) Sarg. 61, 88, 113, 247,
Tsuga (Endl.) Carrière 17, 39, 49, 50, 266, 267, 444, 277, 493, 581, 600, 698, 715, 949, 967, 973, 993,
445, 533, 578, 924, 932, 997, 1007 1007, 1008, 1015, 1016, 1017, 1018, 1029
Tsuga argyrophylla (Chun & Kuang) de Laub. & Tsuga japonica Shiras. 929
Silba 252 Tsuga ×jeffreyi (A. Henry) A. Henry 1007, 1018
Tsuga blaringhemii Flous 1012 Tsuga leptophylla Hand.-Mazz. 1013
Tsuga canadensis (L.) Carrière 62, 597, 752, 779, Tsuga longibracteata W. C. Cheng 533, 1007
974, 991, 1007, 1008, 1009, 1011 Tsuga mairei Lemée & Lév. 981
Tsuga caroliniana Engelm. 87, 1007, 1008, 1009, Tsuga mertensiana (Bong.) Carrière 61, 97, 113,
1010, 1011 428, 491, 493, 564, 581, 600, 619, 654, 715, 993,
Tsuga chinensis (Franch.) E. Pritz. 64, 70, 75, 79, 1007, 1017, 1018, 1029, 1030, 1071
81, 82, 115, 127, 173, 176, 251, 253, 275, 495, 535, Tsuga mertensiana (Bong.) Carrière subsp. grandi­
562, 587, 595, 596, 858, 934, 955, 967, 975, 1007, cona Farjon 99, 1008, 1018
1010, 1011, 1015, 1071 Tsuga mertensiana (Bong.) Carrière subsp. merten­
Tsuga chinensis (Franch.) E. Pritz. var. chinen­ siana 1008
sis 95, 967, 1011 Tsuga mertensiana (Bong.) Carrière subsp. merten­
Tsuga chinensis (Franch.) E. Pritz. var. daibuensis siana var. jeffreyi (A. Henry) C. K. Schneid.
S. S. Ying 1011 1007, 1018
Tsuga mertensiana (Bong.) Carrière subsp. merten­ Widdringtonia cedarbergensis J. A. Marsh 967,
siana var. mertensiana 967, 1018 1020, 1059
Tsuga oblongisquamata (W. C. Cheng & L. K. Fu) Widdringtonia cupressoides (L.) Endl. 1022
L. K. Fu & Nan Li 1012 Widdringtonia dracomontana Stapf 1022
Tsuga patens Downie 1011 Widdringtonia juniperoides (L.) Endl. 1021
Tsuga pattoniana (Balf.) Sénécl. var. jeffreyi A. Hen- Widdringtonia nodiflora (L.) Powrie 968, 1020,
ry 1007, 1018 1022, 1024, 1025
Tsuga sieboldii Carrière 83, 280, 285, 522, 930, 947, Widdringtonia nodiflora (L.) Powrie var. dracomon­
1005, 1007, 1008, 1018, 1019 tana (Stapf) Silba 1022
Tsuga tchekiangensis Flous 1011 Widdringtonia nodiflora (L.) Powrie var. whytei
Tsuga wardii Downie 1013 (Rendle) Silba 1024 1111
Tsuga yunnanensis (Franch.) E. Pritz. 1013 Widdringtonia schwarzii (Marloth) Mast. 1020,
Tsuga yunnanensis (Franch.) E. Pritz. subsp. dura 1023
(Downie) E. Murray 1013 Widdringtonia whytei Rendle 968, 1020, 1024,
Tumion Raf. ex Greene 998 1025
Tumion taxifolium (Arn.) Greene 998 Wollemia W. G. Jones et al. 16, 19, 33, 44, 1026,
1027
Wollemia nobilis W. G. Jones et al. 16, 968, 1026,
V
1057
Veitchia Lindl. 551
Veitchia japonica Lindl. 551
X
Verataxus J. Nelson 969
Xanthocyparis Farjon & Hiep 36, 46, 47, 290, 291,
1028, 1029, 1032
W
Xanthocyparis nootkatensis (D. Don) Farjon &
Washingtonia Winslow 950, 951 Harder 600, 968, 993, 1017, 1028, 1029, 1030
Washingtonia californica Winslow 950 Xanthocyparis vietnamensis Farjon & Hiep 173,
Wellingtonia Lindl. 950, 951 174, 176, 251, 968, 975, 1028, 1030, 1031, 1053,
Wellingtonia gigantea Lindl. 950, 951 1055
Widdringtonia Endl. 22, 37, 46, 48, 1020

End notes
* The following new combination has been published
in this book:

Pinus taiwanensis Hayata var. fragilissima (Busin-

ský) Farjon

* Many names of species and infraspecific taxa have

been typified in this book.