Flora Malaysia

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Flora Malesiana. Series I: Seed Plants
Article in Taxon · February 2006

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Flora Malesiana, Series I, Volume 17 / Part 1 (2006) 1–152
MORACEAE
genera other than FICUS
(C.C. Berg, E.J.H. Corner† & F.M. Jarrett) 1
FOREWORD
The following treatments of Artocarpus, Hullettia, Parartocarpus, and Prainea are

based on the monograph by Jarrett (1959 –1960) and on the treatments she made for
this Flora in cooperation with Dr. M. Jacobs in the 1970s. These included some new
Artocarpus species described in 1975 and the re-instatement of A. peltata. Artocarpus
lanceifolius subsp. clementis was reduced to the species, in A. nitidus the subspe-
cies borneensis and griffithii were reduced to varieties and the subspecies humilis and
lingnanensis included in var. nitidus. The varieties of A. vrieseanus were no longer
recognised. More new Malesian species of Parartocarpus were described by Corner
(1976), Go (1998), and in Artocarpus by Kochummen (1998). A manuscript with the
treatment of the other genera was submitted by Corner in 1972. Numerous changes to
the taxonomy, descriptions, and keys have been made to the original manuscripts, for
which the present first author is fully responsible.
References: Corner, E.J.H., A new species of Parartocarpus Baillon (Moraceae). Gard. Bull. Singa-
pore 28 (1976) 183 –190. — Go, R., A new species of Parartocarpus (Moraceae) from Sabah. Sandaka-
nia 12 (1998) 1– 5. — Jarrett, F.M., Studies in Artocarpus and allied genera I–V. J. Arnold Arbor. 50
(1959) 1– 37, 113 –155, 298 – 368; 51 (1960) 73 –140, 320 – 340. — Jarrett, F.M., Four new Artocarpus
species from Indo-Malesia (Moraceae). Blumea 22 (1975) 409 – 410. — Kochummen, K.M., New
species and varieties of Moraceae from Malaysia. Gard. Bull. Singapore 50 (1998) 197– 219.
MORACEAE
Moraceae Link, Handbuch 2 (1831) 444 (Moriformes), nom. cons.; Baill., Hist. Pl. 6
(1875) 137– 216 (Ulmacées); Benth. & Hook.f., Gen. Pl. 3, 1 (1880) 341– 395 (Urti-
caceae); Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 66 – 98; Corner, Gard.
Bull. Singapore 19 (1962) 187– 252; C.C. Berg, Taxon 27 (1978) 39 – 44; Proc. Kon.
Ned. Akad. Wetensch. C, 91 (1988) 345–362; Cronquist, An integrated system of
classification of flowering plants (1981) 195 –199.
Trees, shrubs, woody climbers, or herbs (Dorstenia, Fatoua), terrestrial, hemi-epi-
phytic (or holo-epiphytic), dioecious or monoecious, with milky sap. Leaves spirally
arranged or distichous, (sub)opposite or subverticillate; stipules fully amplexicaul or
semi-amplexicaul and lateral or intrapetiolar, free or connate; lamina basally attached
(or peltate), with the margin entire or incised (to seemingly compound), venation pin-
nate or subpalmate, brochidodromous. Inflorescences typically in pairs, unisexual or
bisexual, racemose, spicate, globose-capitate, capitate with a discoid to cup-shaped
1) With contributions by P. Baas (wood anatomy), R.P.W. Kromhout & R.W.J.M. van der Ham (pollen
morphology), R. Hegnauer (phytochemistry and chemotaxonomy). Most of the original drawings
are by R. van Crevel and some by E.J.H. Corner.
Flora Malesiana, Ser. I, Vol. 17 / Part 1 (2006)
receptacle (and then with or without involucre), or with an urceolate receptacle, multi-
to uniflorous, bracteate. Flowers unisexual, free or connate (or also adnate to the re-
ceptacle). Staminate flowers: tepals 2 – 4(–7) and free or connate, or perianth lacking;
stamens 1– 4(– 6), straight or inflexed before anthesis; pistillode present or absent.
Pistillate flowers: tepals (3 –)4(– 8), free or connate; pistil 1, ovary unilocular, free
or adnate to the perianth; stigmas 1 or 2, various in shape; ovule 1, (sub)apically at-
tached, anatropous to campylotropous. Fruit an achene or drupaceous (dehiscent or
indehiscent), free or adnate to the perianth, often forming a drupaceous whole with the
fruiting perianth or also with the (fleshy) receptacle. Seed large without endosperm or
small with endosperm; embryo various.
DISTRIBUTION
The family comprises 37 genera and c. 1050 species represented by 14 native genera
with in total 422 indigenous species; 8 species are introduced in Malesia of which
3 belong to Ficus.
The family is essentially tropical. Two genera, Broussonetia and Morus, are associ-
ated with and morphologically adapted to northern warm-temperate conditions. The
genera Ficus and Maclura extend with a few members into warm-temperate conditions
in the northern hemisphere and with two species of Streblus into those of the southern
hemisphere.
Most species of Moraceae (c. 600 spp.) occur in the forest complex of tropical
Asia and Australasia, where the large number of Ficus and, to a lesser extent, of Ar
tocarpus swell the total. Africa with c. 185 species and America with c. 270 species
are specifically poorer, but are richer in endemic genera: 7 out of 17 and 14 out of 19,
respectively.
The third largest genus of the Moraceae, Artocarpus, is largely Malesian. About half
of the species of the largest genus of the family, Ficus, occurs in the Malesian region.
The Asian element links closely to the Australasian, which can be regarded as an
extension of the Asian. It is distinguished by the monotypic genus Sparattosyce, en-
demic to New Caledonia, and by some endemic or subendemic subdivisions of Ficus
and Streblus.
Only six of the native genera, apart from Sparattosyce, are confined to the Asian –
Australasian region. The other genera also occur in Madagascar, the African continent,
and/or the Neotropics.
There are several links between the Asian moraceous flora and the African one.
One pattern is formed by a number of subdivisions of Ficus (subsect. Pedunculatae of
subg. Pharmacosycea, subsect. Urostigma of subg. Urostigma, sect. Sycidium of subg.
Sycidium, and subsect. Ficus of subg. Ficus) which are centred in western Asian dry or
seasonal forest but with linking elements in African savannah woodland. Another pat-
tern is that between the Madagascan region, the eastern Asian mainland, and Malesia
exhibited by the genera Bleekrodea, Broussonetia, Fatoua, Streblus, Trophis, and Ficus
subg. Sycomorus sect. Sycomorus. Other links are formed by the close relationship
between the African genus Treculia and the Asian genus Artocarpus and the African
genus Milicia and Asian Moreae, and by Antiaris toxicaria ranging from West Africa
Berg, Corner & Jarrett — Moraceae (genera other than Ficus)
to the Tonga Islands. Two other species occur in Asia and Africa: Ficus exasperata and
F. palmata.
The links with America are much weaker and are realized by Ficus (subg. Pharma
cosycea) and Trophis, which connect the north western part of the Neotropics to the
eastern part of the Asian –Australasian region. The Neotropics and Africa are linked by
the tribes Dorsteniae and Castilleae.
Ficus, Maclura, and Trophis are pantropical genera. The essentially northern warm-
temperate genus Morus extends into the montane tropics of Asia and America and
lowland Africa. Dorstenia, speciose in America and Africa, is represented in Asia by
only one Indian species.
Within Malesia the greatest numbers of Moraceae coincide with those of the Dip-
terocarpaceae, except for the development of Ficus in New Guinea. Most genera (and
subgenera of Ficus) have one or more widespread species. A Sino-Himalayan ele-
ment infiltrates the Malay Peninsula and some of the species reach Sumatra, Java, and
Borneo. An Australian element is evident in Ficus subsect. Malvanthera. It penetrates
only the adjacent part of the Malesian region and the Pacific. The Australasian Streblus
sect. Parastreblus extends to New Zealand and New Caledonia. The genera endemic to
Malesia are Antiaropsis, restricted to New Guinea, and the more widespread Prainea.
Parartocarpus slightly exceeds Malesia.
Two neotropical genera, Castilla and Dorstenia, have been introduced, each with one
species. A few other species introduced in the Malesian region belong to Broussonetia,
Ficus, and Morus and are of Asian origin.
FIELD CHARACTERS
The presence of milky sap (latex) and stipules which often leave circular scars distin-
guish Moraceae. The former feature is shared with Euphorbiaceae, for which diagnostic
features are the geniculate petiole and the presence of glands at the apex of the petiole,
or on the lower margin of the lamina. In the majority of the Ficus species, waxy glands
occur on the lower surface of the lamina, often in the axils of the basal lateral veins,
or on the base of the midrib. Such glands are absent in other Moraceae. If not by the
waxy glands, the majority of the Ficus species can be distinguished by aerial secondary
roots that make them hemi-epiphytes (‘stranglers’), or root climbers, whereas many
others are cauli- or flagelliflorous. Smooth grey bark, marked with leaf scars and exud-
ing latex on cutting, will also distinguish moraceous trees. A bright orange-brown or
grey-brown roughish bark, with thick copious latex, will often distinguish Artocarpus.
Dark red lateral roots with latex will distinguish some species of Artocarpus, as yellow
roots with latex will distinguish Parartocarpus. The presence of thorns distinguishes
the climbers of Maclura and the small trees of several Streblus species. Antiaris shows
the architectural ‘model of Cook’ (Hallé & Oldeman 1970; Berg 1977; Hallé et al.
1978), characterised by spirally arranged leaves on the trunk and main branches, and
distichous on the horizontal lateral branches which also may bear the inflorescences.
Each axillary bud produces a lateral branch which ultimately is detached (by abscission)
leaving a concave scar.
References: Berg, C.C., The Castilleae, a tribe of the Moraceae, renamed and redefined due to the
exclusion of the type genus Olmedia from the ‘Olmedieae’. Acta Bot. Neerl. 26 (1977) 73 – 82. — Hallé,
F. & R.A.A. Oldeman, Essai sur l’architecture et la dynamique de croissance des arbres tropicaux.
Masson, Paris (1970). — Hallé, F., R.A.A. Oldeman & P. B. Tomlinson, Tropical trees and forests – an
architectural analysis. Springer-Verlag Berlin, Heidelberg, New York (1978).
MORPHOLOGY
Habit — The family consists chiefly of lowland trees. In Malesia, tall canopy trees or
emergents are found in Antiaris, Artocarpus, Ficus, Parartocarpus, Prainea, and Stre
blus. They often have buttresses. Smaller, undergrowth trees of varying stature belong
to Antiaropsis, Broussonetia, Ficus, Hullettia, Streblus, and Trophis. Shrubs and treelets
are found in Bleekrodea, Ficus, Maclura, and Streblus. Species which can become tall
trees, such as Streblus elongatus, may flower as shrubs or treelets.
Most tree species are evergreen, but some are deciduous both in seasonal and ever-
wet forest. The trees have intermittent growth, with sometimes conspicuous buds that
are temporarily dormant.
Monocaulous or sparingly branched trees, with relatively thick branches and terminal
tufts of large leaves (pachycladous, or in Corner’s terminology pachycaul, trees), occur
in several species of Ficus (e.g., F. pseudopalma), one species of Dorstenia (Africa),
and one species of Naucleopsis (America). A wide range of tree forms occur between
the pachycladous tree and the more common leptocladous one, with slender branches
and small leaves mostly evenly distributed on the leafy twigs.
There are c. 100 species of climbers in Malesia, many of them belonging to Ficus and
Maclura. Moreover, this habit is found in single species in Broussonetia and Trophis,
and sometimes in Prainea scandens. The climbing habit is rare in Africa and America,
in each restricted to a single species of Maclura and to one or two (sub)lianescent Ficus
species. The climbers may be twining or straggling (Trophis scandens and species of
Ficus subg. Sycidium sect. Palaeomorphe), scramblers with thorns (Maclura), or root
climbers (Ficus subg. Synoecia).
Most species are unarmed, although thorns are found in Maclura and Streblus, in the
former often to assist climbing. Spiny leaf margins and/or apices occur in some species
of Streblus and also in the neotropical genera Clarisia and Sorocea.
Fatoua is predominantly herbaceous as are the majority of Dorstenia. In the latter
genus there are a wide range of herbaceous life forms: stem succulents, tuber succulents,
geophytes, epiphytes, and annuals (Berg & Hijman 1999). In both of these genera, plants
can be suffrutescent. Suffrutescence is also known in the essentially woody Ficus (e.g.,
F. griffithii Miq. (Asia) and F. suffruticosa Corner (New Guinea)) and the neotropical
Perebea (P. humilis C.C. Berg).
Hemi-epiphytism is a prominent life form in Ficus, which also has two species that
can be holo-epiphytes. The very wide range of life and growth forms in Ficus are de-
scribed under the genus.
The twig apices are shed in Morus and Broussonetia, and elongation of twigs is
performed by the meristem in the upper most lateral bud; the resting buds are scaled.
This trait is also found in other elements of northern temperate forest, such as Tilia and
Ulmus. Shoot apices are even abscised in tropical lowland species of the two moraceous
genera.
Trichomes — The unicellular hairs vary in length, diameter, shape, septation, and col-
our; they show particularly great variation in Ficus, including interwoven and irritating
hairs. Uncinate hairs are quite common as in most other Urticales. They occur frequent-
ly in the Dorstenieae, often in Artocarpeae and Moreae, but are absent in Castilleae,
and in Ficus only known from three species. Cystolith hairs form scabrous surfaces
as in many Ficus species and Antiaris (Renner 1907). Pluricellular trichomes are very
common and varied in shape and colour. The pluricellular, globose or ellipsoid, capitate
trichomes that occur in (nearly?) all species of Moraceae have often been described as
glandular.
They are mostly whitish and can be found on the young plant parts. The heads vary
in shape (Bhat & Kachroo 1979) and may have diagnostic value. Pluricellular trichomes
may also be elongate and can be found in many species of Ficus (in particular of the
monoecious subgenera), and are then mostly brown.
Exudates — The family is characterized by the presence of latex, usually milky, some-
times ± watery (as in some Ficus species) or entirely watery as in Fatoua. The latex
can be poisonous as in Antiaris toxicaria, Ficus magnoliifolia, and Parartocarpus
venenosus, or coagulate into rubber (as in Ficus elastica). That of Ficus tinctoria is
used to prepare a red dye (Florence 1997).
Glands — Nearly all species of Ficus have glandular spots with a waxy surface. They
occur on the leaves or on the nodes of leafy twigs (see FM 17-2: 84). Glands in similar
positions to many Ficus species are found in most species of Artocarpus subg. Pseudo
jaca (see p. 108); it is not clear whether they exude wax as in Ficus. Artocarpus alt
issima has glands on the margin of the lamina, which is unique in Moraceae.
Leaves — The leaves are mostly alternate and spirally arranged or distichous. Opposite
leaves are found in some groups of Ficus (subg. Sycidium and subg. Sycomorus), often
in combination with alternate arrangement. Opposite leaves may also occur in Brous
sonetia papyrifera and are characteristic for the neotropical genus Bagassa. Subverti-
cillate leaves occur in some species of Ficus subg. Sycidium. Such arrangement of the
leaves occurs occasionally in Artocarpus (subg. Pseudojaca).
The lamina varies from large, up to 2 m long in Ficus solomonensis or up to 1 m
wide in F. dammaropsis, to very small, not more than 3 cm long in F. vaccinioides
King from Taiwan, and from 2 cm in F. humbertii C.C. Berg from Madagascar, or some
neotropical Dorstenia species. The lamina is usually basally attached (peltate in some
species of Dorstenia). It is mostly entire, less commonly pinnately or palmately incised.
Pinnately incised laminas occur in some species of Artocarpus, Dorstenia, and Ficus,
palmately incised ones in Broussonetia, Dorstenia, Ficus, and Morus. The venation is
mostly brochidodromous. The tertiary venation is basically scalariform, with numer-
ous parallel transverse veins in the intercostal area. This type of venation evolved into
a reticulate pattern, and, in some groups of Ficus, subsequently into tertiary venation
largely parallel to the secondary venation.
The texture of the lamina varies from thickly coriaceous to chartaceous, or even
membranaceous in herbaceous genera. The margin is usually entire in coriaceous lami-
nas (of evergreen forest species) and often dentate in thinner ones. The petiole varies
from long to short. Long petioles are generally associated with cordiform laminas.
The stipules are often fully amplexicaul, leaving annular scars, by which so many
Moraceae can be recognized, but they may also be semi-amplexicaul or lateral. The
stipules vary from large (more than 10 cm long) to small (down to 1 mm long). They
are mostly caducous but may be persistent or subpersistent.
Inflorescences — The inflorescences are mostly paired as in other families of the Ur-
ticales. The types of inflorescences range from cymes, to racemes, to spikes (slender
or thick and almost spadix-like), to heads, either globose- to ellipsoid-capitate, or
discoid (bisexual or unisexual, involucrate or not), or to urceolate structures (opening
or remaining closed).
They have a simple structure: racemose, spicate, cymose, or capitate, and mostly
unisexual in the tribe Moreae and in the neotropical genera of the tribe Artocarpeae.
The staminate ones often resemble catkins of Amentiferae, but the pistillate ones in fruit
look different owing to the fleshy pericarp, perianths and/or interfloral bracts. In the
majority of the Moraceae the inflorescences are complicated by condensation of axes,
by fusion of homologous or adjacent structures, more prominent bracts, and bisexual-
ity of inflorescences. These complex structures are mostly the functional entities with
regard to pollination, and in fruiting state, to dispersal, as is very clearly the case in
the syconium or fig of Ficus. The complex inflorescences can be pseudoflorous, but
are more often pseudocarpous and they usually bear numerous flowers. Reduction to
the uniflorous state sometimes occurs (e.g., in Ficus oleifolia, and in the neotropical
Perebea humilis).
The abaxial sterile strip or groove, which is particularly clear in elongate inflores-
cences, indicates adaxial orientation of the flowers in essentially racemosely construct-
ed inflorescences. Pistillate flowers tend to occur in the centre of bisexual (essentially
cymosely constructed) inflorescences and staminate flowers in the periphery (Berg
1977).
Some flowers and inflorescences show clear adaptation to the mode of pollination.
In other cases the primary modifications seem to be for the protection of developing
stamens and ovaries against predation by insect larvae (cf. Berg 1990). This protection
can often be related to insects breeding in staminate inflorescences before later pollina-
tion.
The inflorescences are mostly axillary. They may occur below the leaves on previous
season’s growth, often so in deciduous species. Ramiflorous inflorescences are born
on short-shoots which are often already present in the leaf axils and often continue to
bear inflorescences below the leaves, even down to the smaller branches. Cauliflorous
inflorescences on short spur-like branchlets or on branches with long internodes are
found in Artocarpus and in Ficus. In the latter genus such branches produced at the
base of the trunk may become stolon-like (flagelliflory or geocarpy). Cauliflory is also
found in the African genus Treculia and the neotropical genus Clarisia.
Flowers — Moraceous flowers are small, monochlamydeous, and usually unisexual.

Flowers with stamens and non-functional pistils are found in Ficus subg. Sycidium.
The flower is often 4-merous, but the number of tepals may be more or less than 4, or
the perianth may be absent (e.g. in Brosimum (male, America), in Hullettia (male and
female?), Treculia (female, Africa), and Trilepisium (male, Africa), or strongly reduced
as in several species of Ficus subg. Sycomorus sect. Sycocarpus (female, Asia).
The number of stamens is rarely more than 4 (e.g. in subg. Ficus), but often less;
Artocarpus and many species of Ficus have only one. The tepals are either imbricate
or valvate (as in Trophis), or narrow and not touching or overlapping each other (e.g. in
Ficus subg. Synoecia). They may be free, or connate, forming tubular perianths, which
can be fused with the ovary (e.g. in Trophis). Perianths of pistillate flowers either are
or become ± fleshy; Fatoua and Ficus are notable exceptions.
The ovaries contain a single apically or subapically attached, anatropous to campylo-
tropous ovule. There are two stigmata, or one by reduction. The stigmata vary in shape
from filiform, to tongue-shaped, to truncate. Heterostyly is found in Ficus.
The stamens are straight in the bud in most genera, but in the majority of the species
of the tribe Moreae inflexed and at anthesis bend outwards elastically. The anthers have
two thecae but, due to fusion, seemingly only one in Ficus subsect. Malvanthera.
Pistillodes are found in flowers with the urticaceous type of stamens where they may
play a role in keeping the anther in position. They are, moreover, characteristic for Ficus
subg. Sycidium, in which they may be as large as pistils. Pistillodes are often found in
flowers of Ficus, e.g. in subg. Pharmacosycea and only occasionally in other groups
of Moraceae.
Sexuality — The flowers are unisexual, or in some species of Ficus subg. Sycidium
sect. Palaeomorphe, functionally so. The inflorescences are bisexual or unisexual and
the plants monoecious or dioecious.
Bisexual inflorescences occur in Bleekrodea, Fatoua, Ficus (subg. Pharmacosycea,
subg. Urostigma, and some species of subg. Sycomorus), in some Streblus species, as
in S. elongatus, S. tonkinensis (Eberh. & Dubard) Corner, and S. asper (sometimes),
and in most of the Dorstenieae. Bisexuality of inflorescences is usually found in those
that are basically cymose and circular in outline.
Monoecism, combined with unisexual inflorescences, is found in Artocarpus, Hul
lettia, and Streblus p.p. A single pistillate inflorescence with several staminate inflores-
cences on the same node (or short-shoot) are often found in species of the Castilleae,
as in Antiaris and Castilla.
Dioecism is found in Antiaropsis, Broussonetia, Maclura, Morus, Parartocarpus,
Prainea, Streblus p.p., and Trophis. In Ficus, the subgenera Ficus, Sycidium, Synoecia,
and Sycomorus p.p. are morphologically gynodioecious, but functionally dioecious. In
functionally female inflorescences of Ficus, neuter flowers may substitute the staminate
flowers.
Androdioecism, in combination with bisexual and unisexual inflorescences, is known
from the American Helianthostylis Baill. It may occur in the African Bosqueiopsis De
Wild. & T. Durand, and in combination with unisexual inflorescences in Castilla.
Fruits — The distribution of the family suggests that the dehiscent drupe might be the
basic fruit type of Moraceae (Berg 1977). The exocarp is whitish, stipitate at the base,
and laterally thickened. The turgid halves clasp the endocarp body, kept in its position
by the vascular bundle running on top of it. When this bundle breaks the endocarp
body is ejected from the fruit if it is small, or is squeezed out if it is large. It can also
be ejected as in Streblus elongates and S. macrophylla, in the latter up to 10 m from
the tree (pers. comm. Dr. D.J. Middleton). Small endocarp bodies, as can be found in
Fatoua (and Dorstenia), are white and tuberculate; large ones are black and smooth
(Antiaropsis, Streblus). Dehiscent drupes are found in Antiaropsis, Bleekrodea, Brous
sonetia, Dorstenia (except for the only epiphytic species, see Berg & Hijman 1999),
some species of Ficus, Fatoua, Scyphosyce Baill. (Africa), Sparattosyce Bureau (New
Caledonia), and Utsetela Pellegr. (Africa). Some of the features of the pericarp of the
dehiscent drupe, such as the basally narrowed and/or laterally thicker pericarp, can also
be found in indehiscent types.
The ovary is often fused with the perianth and the features described above are
obliterated. Fruits with dry pericarps are found in the majority of the Ficus species. The
stipitate base often found in the achene of Ficus might be homologous to the stipitate
base of the dehiscent fruit.
The seed coat, protected by the more or less hard endocarp, is thin, but often conspicu-
ously vascularised. Small seeds contain endosperm, as in Ficus, but in large seeds it is
(almost) lacking. The embryos of small seeds are simple, those of large seeds more elabo-
rate with thick, folded and/or unequally large cotyledons and short or long radicles.
References: Berg, C.C., Urticales, their differentiation and systematic position, in: K. Kubitzki (ed.),
Flowering plants – Evolution and classification of higher categories. Pl. Syst. Evol., Suppl. 1 (1977)
349 – 374. — Berg, C.C., Differentiation of flowers and inflorescences of Urticales in relation to their
protection against breeding insects and to pollination. Sommerfeltia 11 (1990) 13 – 34. — Berg, C.C.
& M.E. E. Hijman, The genus Dorstenia (Moraceae). Ilicifolia 2 (1999) 1– 211. — Bhat, M.M. & P.
Kachroo, The trichome in phylogeny of Urticales, in: C.P. Malik (ed.), Annual Rev. Pl. Sci. 1 (1979)
375 – 407. — Florence, J., Flore de la Polynésie Française 1. Paris (1997). — Renner, O., Beiträge zur
Anatomie und Systematik der Artocarpeen und Conocephaleen inbesondere der Gattung Ficus. Bot.
Jahrb. Syst. 39 (1907) 319 – 448.
WOOD ANATOMY
(P. Baas - Leiden & E.A. Wheeler - Raleigh, North Carolina)
Wood anatomy — For a very full bibliography of the wood anatomy of the Moraceae
see Gregory 1994. The microscopic wood structure of numerous Malesian taxa is also
documented with coded wood descriptions (following the standard codes of the IAWA
Committee 1989) on the InsideWood web page (http://insidewood.lib.ncsu.edu). The
general description of the Malesian genera treated in this volume (i.e., Moraceae exclud-
ing Ficus) is based on these coded descriptions, which in turn are based on literature
cited at the end of this section. Of the genera Bleekrodea, Fatoua, Hullettia no wood
anatomical data were available to us.
Heartwood is either distinct and dark, or not clearly demarcated from the sapwood.
The texture of the wood is usually medium to coarse. Interlocked grain fairly often oc-
curs in the family.
Growth ring boundaries are absent or faint except in some species of Antiaris, Arto
carpus, Broussonetia, Maclura, Morus, and Prainea where they can be distinct. Wood
typically diffuse-porous, but semi-ring-porous to ring-porous in Broussonetia p.p.,
Maclura, and Morus p.p, rarely with a tendency towards tangential (Morus) or diagonal
to dendritic (Antiaris p.p.) vessel pattern.
Vessels both solitary and in short radial multiples; in Artocarpus p.p., Maclura,
and Morus also partly in clusters. Vessel diameter usually ranging from 100 – 200 µm
but smaller in some species, wider in others; vessels tending to be of two size classes
(with the smaller vessels in clusters or multiples) in Antiaris, Broussonetia, and Morus.
Vessel frequency usually low (< 5/ mm2) to intermediate (5 – 20 mm2); more rarely
higher, especially in Streblus p.p. Perforations simple. Intervessel pits non-vestured,
alternate, round to polygonal, medium-sized to large (>> 7 µm diam.), but small (4 –7
µm) in Broussonetia p.p., Dorstenia, Prainea p.p., and Streblus p.p. Vessel-ray pits typi-
cally with reduced borders to simple, elongate or round; non-reduced pit borders occur
in Artocarpus p.p., Morus p.p., Prainea p.p., and Trophis. Tyloses commonly present in
heartwood. Gummy and other deposits only recorded in Artocarpus p.p. Helical vessel
wall thickenings present in Maclura (variable) and Morus.
Fibres libriform (that is, with simple to minutely bordered pits largely confined to
the radial walls); mostly non-septate, but septate in Antiaris, Antiaropsis, Castilla, and
Prainea; ranging from thin-walled to very thick-walled in close correlation with the
density and hardness of the timber.
Parenchyma predominantly paratracheal, usually in a combination of scanty para
tracheal to vasicentric and aliform to confluent. Regular tangential parenchyma bands
occur in Artocarpus p.p., Dorstenia p.p., Maclura, Prainea p.p., Streblus, and Trophis.
Marginal parenchyma bands may occur in species with growth ring boundaries. Paren-
chyma strands 2 – 4(– 8) cells long.
Rays varying in width and mostly including broad rays over 4 cells wide, but multi-
seriate rays 2- or 3-seriate in Antiaropsis, Broussonetia p.p., Dorstenia, Parartocarpus
p.p., Streblus p.p., and Trophis; typically heterocellular with 1– 4 (or even more) mar-
ginal rows of square to upright cells; homocellular and composed of procumbent cells
only in Parartocarpus and Maclura p.p. Sheath cells of varying distinctness present in
some of the taxa.
Laticifers usually present in the rays, absent from Artocarpus p.p., Morus, Prainea,
and Streblus. Prismatic crystals recorded in rays and axial parenchyma cells of Antiaris,
Broussonetia, Maclura, Morus, Parartocarpus p.p., and Streblus p.p. Druses recorded
in Broussonetia kurzii. Silica bodies absent, but vitreous silica sometimes present in
some of the genera.
The wood anatomical diversity in the Moraceae is of great systematic significance and
also allows microscopic wood identification down to the genus or to groups of related
genera. A detailed comparison of the wood diversity pattern in the Moraceae from all
over the world, as comprehensively documented in a series of papers by Koek-Noor-
man, Ter Welle, and Topper with a modern DNA-based phylogenetic classification of
the Moraceae is long overdue.
10 Flora Malesiana, Ser. I, Vol. 17 / Part 1 (2006)
Literature (selected references only; for general references on the wood of Malesian Moraceae and
a comparison with the wood anatomy of Ficus, see Flora Malesiana Ser. I, 17/2 (2005) 9 –11).
Gregory, M., Bibliography of systematic wood anatomy of Dicotyledons. IAWA J. Suppl. 1 (1994).
— IAWA Committee, IAWA list of microscopic features for hardwood identification. IAWA Bull.
n.s. 10 (1989) 219 – 332. — Koek-Noorman, J., S.M.C. Topper & B.J.H. ter Welle, The systematic
wood anatomy of the Moraceae (Urticales). I. Tribe Castilleae. IAWA Bull. n.s. 5 (1984a) 183 –195.
— Koek-Noorman, J., S.M.C. Topper & B.J.H. ter Welle, The systematic wood anatomy of the Mora-
ceae (Urticales). II. Tribe Dorstenieae. IAWA Bull. n.s. 5 (1984b) 317– 329. — Kribs, D.A., Com-
mercial foreign woods on the American market. Dover Publications, New York (1968). — Lemmens,
R.H.M.J., I. Soerianegara & W.C. Wong (eds.), Plant Resources of South-East Asia 5 (2). Timber
trees: Minor commercial timbers. Backhuys Publishers, Leiden (1995). — Metcalfe, C.R. & L. Chalk,
Anatomy of the Dicotyledons. Clarendon Press, Oxford (1950).— Normand, D. & J. Paquis, Manuel
d’identification des bois commerciaux. Tome 2. Afrique guineo-congolaise. Centre Technique Fores-
tier Tropical, Nogent s/Marne (1976). — Pearson, R.S. & H.P. Brown, Commercial timbers of India.
Central Publ. Branch, Govt. of India, Calcutta (1932).— Purkayastha, S.K., K.B.S. Juneja & S.M.H.
Kazmi, Anatomy of the more important Andaman commercial timbers (with notes on their supply,
properties, and uses). Indian For. Rec. (N.S. Wood Anatomy) 2 (1) (1976) 1– 48. — Sosef, M.S.M.,
L.T. Hong & S. Prawirohatmodjo (eds.), Plant Resources of South-East Asia 5 (3). Timber trees:
Lesser-known timbers. Backhuys Publishers, Leiden (1998). — Ter Welle, B.J.H., J. Koek-Noorman
& S.M.C. Topper, The systematic wood anatomy of the Moraceae (Urticales). IV. Genera of the tribe
Moreae with urticaceous stamens. IAWA Bull. n.s. 7 (1986a) 91–128.— Ter Welle, B.J.H., J. Koek-
Noorman & S.M.C. Topper, The systematic wood anatomy of the Moraceae (Urticales). V. Genera of
the tribe Moreae without urticaceous stamens. IAWA Bull. n.s. 7 (1986b) 175 –193.
POLLEN MORPHOLOGY
(R.P.W. Kromhout & R.W.J.M. van der Ham)
Pollen descriptions and illustrations of 25 out of the 37 genera of Moraceae are available
(literature cited by Thanikaimoni & Van der Ham 1999). Of the current seven tribes,
Antiaropsideae, Artocarpeae, Castilleae, Dorstenieae, Ficeae, Moreae, and Soroceae
(Datwyler & Weiblen 2004, Berg 2005), the tribes Antiaropsideae, Artocarpeae, Cas-
tilleae, Moreae, and Soroceae show a similar pollen type. The pollen grains belonging
to this type are very small to medium-sized (9 – 33 µm) and 2 – 6-porate. The shape of
2-porate grains is ellipsoid to spheroidal, if ellipsoid, then often slightly asymmetri-
cal. The shape of 3-porate grains is spheroidal to almost triangular in polar view and
oblate to spheroidal in equatorial view. Pollen grains with more than three pores are
pantoporate (never stephanoporate) and suboblate to spheroidal. The number of pores
per pollen grain may vary within a species and even within individuals. The pores are
circular and usually 1– 2.5 µm in diameter, but a few species of Morus show pores up to
5.5 µm. The exine is thin, up to 1.5 µm thick and tectate. The ornamentation is clearly
scabrate using scanning electron microscopy, but indistinctly scabrate or psilate using
light microscopy. Commonly, the sexine and nexine are more or less equally thick, while
the infratectum is indistinctly columellate. Sometimes, exine stratification is indistinct,
using light microscopy. The pollen of tribe Ficeae (Ficus) is similar, but smaller (7– 22
µm) and 2- or sometimes 3-porate (Langeveld & Van der Ham 2005). Tribe Dorstenieae
has different pollen, although the size range is more or less the same (10 – 33 µm) and
grains with more than three pores are also pantoporate. The sexine is always thicker
than the nexine, and nearly always only faintly stratified. The pollen of Brosimum,
Berg, Corner & Jarrett — Moraceae (genera other than Ficus) 11
Helianthostylis, and Trymatococcus is much like the 2-porate pollen of the previous
type, but Brosimum has a thicker, undifferentiated exine, and the pores may have an
annulus and an operculum. Helianthostylis and Trymatococcus have an annulus and no
operculum. The pollen grains of the monotypic genus Bosqueiopsis are 3- or 4-angular
or asymmetrical and 22 – 27 µm in diameter. They are 3 – 6-porate, with slightly annu-
late pores up to 6 µm. The exine is thin, flexible and scabrate. The genus Trilepisium
has pollen like that of Bosqueiopsis, but the grains are smaller (19 – 23 µm), have more
(8 –12) and slightly smaller pores without annuli or aspides. The scabrate ornamentation
is slightly more coarse. Utsetela, another monotypic genus, has according to Hoen &
Punt (1989) the most primitive pollen in the tribe. It is pantoporate, with 15 – 20 sunken
pores, and a thick, slightly scabrate exine with distinct columellae. It lacks annuli, as-
pides and operculi. Dorstenia is the largest genus in the tribe (c. 170 species). Hoen &
Punt (1989) found nine different pollen types in a study of approximately one third of
the total number of species. Dorstenia pollen is ellipsoid to spheroidal. The grains are
6 – 80-pantoporate, and may have annuli, aspides, and operculi.
The pores are usually distinctly sunken. The exine is mostly thin, but varies in
thickness due to the presence of annuli, aspides, and the often verrucate ornamenta-
tion. Usually, scabrae are present as well. Columellae are faint, but visible, using light
microscopy.
An important factor in pollen morphological diversity in plant groups is the presence
of different pollination syndromes. Unfortunately, pollination data of Moraceae (except
Ficus) are scarce. Very little is known of pollination in the pollen morphologically most
diverse tribe Dorstenieae. Beetles have been found on flowers of Dorstenia species,
but their role in pollination has not yet been satisfactorily demonstrated (Hoen & Punt
1989, Berg & Hijman 1999). Other Dorstenia species have flower types that are usually
associated with wind pollination, or have flowers that seem to suggest pollination by
flies (Berg & Hijman 1999). Further pollination studies may elucidate the origin of the
remarkably diverse pollen morphology in the Dorstenieae.
References: Berg, C.C., Blumea 50 (2005) 535–550. — Berg, C.C. & M. Hijman, Ilicifolia 2 (1999).
— Datwyler, S.L. & G.D. Weiblen, Amer. J. Bot. 91 (2004) 767–777. — Hoen, P.P. & W. Punt, Rev.
Palaeobot. Palynol. 57 (1989) 187– 220. — Langeveld, J.M. & R.W.J.M. van der Ham, Flora Malesi-
ana, Ser. I, 17, part 2 (2005) 50 – 51. — Thanikaimoni, K. & R.W.J.M. van der Ham, Publ. Dép. Ecol.
Inst. Fr. Pondichéry 39 (1999).
PHYTOCHEMISTRY AND CHEMOTAXONOMY

(R. Hegnauer)
Relatively few members of the family have been examined thoroughly by phytochem-
ists. So the chemical characterisation must be regarded as a very incomplete and provi-
sional one. The following features seem to be typical to some extent of Moraceae:
1) Most members of the family deposit large amounts of silicic acid and calcium car-
bonate in the walls of leaf cells, especially hairs and unspecialised epidermal cells.
Cystoliths and crystals of calcium oxalate are ubiquitous.
2) Mucilage cells and ducts occur frequently. Thorough chemical characterisation of
different kind of mucilage of Moraceae are still lacking.
3) Latex-cells and non-articulated latex tubes are widespread. Some species produce
rubber-rich latex. In other species, ‘resins’ (triterpenic alcohols and their acetates
and cinnamates) or ‘waxes’ (ethers of cerotinic acid, esters of triterpenic alco-
hols with several fatty acids) predominate. Still other species produce protein-rich
latex; ficin is a papain-like enzyme from the latex of several species of Ficus, ficin
containing latex is used locally as anthelmintic (in the Neotropics).
A few genera are known to contain cardio-toxic compounds; these have been
isolated from, or demonstrated to be present in, latex of Antiaris, Castilla, Naucle
opsis (= Ogcodeia), barks of Streblus, and seeds of Antiaris, Antiaropsis, Castilla,
and Naucleopsis. The latex of Antiaris toxicaria is used as arrow poison in eastern
Asia, and that of some species of Naucleopsis for the same purpose in the Neotrop-
ics (Bisset & Hylands 1977). Still other species seem to produce latexes which are
very rich in phenolic compounds; Vreede (1949) observed flavonoids (not definitely
defined) in yellow latexes of several species of Ficus. Chlorogenic acid was isolated
from the latex of Castilla elastica.
4) Polyphenolic compounds are common in Moraceae. Derivates of p-coumaric acid,
caffeic acid, kaempferol, and quercetin occur frequently in leaves. Myricetin is less
common. Leucoanthocyanins are absent of the leaves of many species, but often
present in species of Ficus. The so-called tannin-idioblasts of the anatomical litera-
ture are present in the mesophyll of many species. They probably contain catechins
and leucoanthocyanins, but no true tannins, which have yet to be demonstrated in
Moraceae for certain.
5) The compounds of most interest from the systematic point of view are the highly
characteristic phenolic compounds which seem to be rather common in the roots,
stems, fruits, and sometimes the leaves. These phenolics may roughly be classified
in four groups: C6 -C 3-compounds (coumarins), represented by furanocoumarins
and dimethylpyranocoumarins); C6 -C 1 -C 6-compounds (benzophenones like ma-
clurin and xanthones); C 6 -C 2 -C 6 -compounds (stilbenes like chlorophorin and
hydroxyresveratrol); C6 -C 3 -C 6-compounds (highly characteristic flavonoids like
cyanomaclurin, morin, artocarpetin, cycloartocarpin, and pomiferin). The resist-
ance of the woods of some members of the family to the attack by fungi, insects,
and termites, and the tinctorial properties of the woods of others, are largely due to
such polyphenolic compounds.
6) Saponins and alkaloids seem to be rather rare in Moraceae. Alkaloids are known
from some species of Ficus. No taxonomic implications are apparent at present
from these occurrences.
To sum up, the accumulation of minerals in leaves (SIO 2, CaCO 3), and the production
of a whole array of unusual phenolic compounds represent the most striking, currently
known, phytochemical features of Moraceae.
With regard to the different mineralizations (including the presence of cystoliths) the
family resembles Cannabidacea and Urticaceae.
Some outstanding features of phenolic compounds of Moraceae indicate similarities
in secondary metabolism with members of the rosalean alliance, especially with Legu
minosae.
These features are: production of furanocoumarins, isoflavones (osajin, pomiferin),

stilbenes, flavonoids with a resorcin-type hydroxylation pattern of the B-ring (cyanoma-
clurin, artocarpetin, morin, etc.), and the frequent attachment of isoprenoid substituents
to aromatic rings (coumarins, stilbenes, and flavonoids).
Isoprenylation also marks moraceous xanthones (alvaxanthone, macluraxanthone),
in which respect the family resembles Guttiferae. Knowledge of the distribution of
all these types of phenolics is still rather scanty. It would be cautious therefore not to
put too much weight on the metabolic similarities mentioned. A more comprehensive
review of chemical characters may be found in Hegnauer (1969).
Volatile compounds produced by inflorescences of Ficus play an important role in
attraction of pollinators. Compounds extracted from receptive syconia of F. carica in-
clude benzyl alcohol, linanool, linanool oxides (furanoid), several aromatic compounds
(cinnamic aldehyde, cinnamic alcohol) and indole1 (Gibernau et al. 1997).
References: Bisset, N.G. & P.J. Hylands, Cardiotonic glycosides from latex of Naucleopsis mello-
barretoi, a dart poison plant from north-west Brazil. Econ. Bot. 31 (1977) 307–311. — Gibernau, M.,
H.R. Buser, J.E. Frey & M. Hossaert-McKey, Volatile compounds from extracts of figs of Ficus carica.
Phytochemistry 46 (1997) 241– 244. — Hegnauer, R., Chemotaxonomie der Pflanzen 5. Birkhäuser
Verlag, Basel & Stuttgart (1969). — Vreede, M.C., Topography of the lactiferous system in the genus
Ficus. Ann. Bot. Gard. Buitenzorg 51 (1949) 125–149.
CYTOTAXONOMY
There are two series, one with the haploid number of 13 and the other with 14 (Fedorov
1969; Oniguma & Tobe 1995). Brosimum, Broussonetia, Streblus p.p., and Ficus belong
to the first series. Ficus are mostly diploids, but some (mainly African species) are or
can be tetraploid, and F. elastica triploid (Ohri & Khoshoo 1987). The genera belong-
ing to the second series are Antiaris, Artocarpus, Castilla, Clarisia, Maclura, Morus,
Pseudolmedia, Scyphosyce (cf. Berg 1977), Streblus p.p., and Trophis. Some Arto-
carpus species are tetraploid (or hexaploid in cultivars of A. altilis) and some Morus
species polyploid, up to 2n = 308. Dorstenia shows much variation in its chromosome
numbers: 2n = 24, 26, 28, 30, 32, 36, 38, 40, 42, 48, 52, and c. 64 (Berg & Hijman
1999). Deviating numbers are reported for Maclura tricuspidata, 2n = 50 (Morawetz
& Samuel 1989) and Naucleopsis guianensis, 2n = 20 (Dmitrieva & Parfenov 1985).
The haploid chromosome number of both Cecropiaceae and Ulmaceae is 14, that of
Urticaceae varies from 7 to 14. The chromosome numbers of the Cannabidaceae devi-
ates from the general pattern in the order.
References: Berg, C.C., Revisions of African Moraceae (excluding Dorstenia, Ficus, Musanga
and Myrianthus). Bull. Jard. Bot. État 47 (1977) 267– 407. — Berg, C.C. & M.E.E. Hijman, The
genus Dorstenia (Moraceae). Ilicifolia 2 (1999) 1– 211. — Dmitrieva, S.A. & V. I. Parfenov, Kariolog-
icheskaja kharakteristika nekotorykhvidov poleznykh rastenij flori Belorussii. Izvestii Akademii Nauk
Belorusskoi SSR: Seriia Biologicheskikh Nauk 6 (1985) 3 – 8. — Fedorov, A.A. (ed.), Chromosome
numbers of flowering plants. Leningrad (1969). — Morawetz, W. & M.R.A. Samuel, Karyological
patterns in Hamamelidae, in: P.R. Crane & S. Blackmore, Evolution, systematics and fossil history
of the Hamamelidae 1: ‘Introduction’ and ‘lower’ Hamamelidae (1989) 129 –154. Clarendon Press,
1) This paragraph is an addition by C.C. Berg.

Oxford. — Oniguma, K. & H. Tobe, Karyomorphology of some Moraceae and Cecropiaceae (Urti-
cales). J. Pl. Research 108 (1995) 313 – 326. — Ohri, D. & T.N. Khoshoo, Nuclear DNA contents in
the genus Ficus (Moraceae). Pl. Syst. Evol. 156 (1987) 1– 4.
ECOLOGY
The majority of the species are elements of lowland, mostly evergreen forest or less
frequently, seasonal forest. Trees of coastal, riparian, and secondary growth occur
mainly in Ficus, but Streblus asper is characteristic of the more seasonal forest of
western Malesia.
Some species of Ficus (F. deltoidea and F. oleifolia) often occur on poor sandy soil
in kerangas forest. Other species of Ficus are associated with calcareous substrates
(F. anastomosans, F. calcarata, F. calcicola, F. subcaudata (?), and the form F. tinctoria
described as F. swinhoei). A form (ecotype) of F. ulmifolia is associated with extreme
ultra basic soil (Philippines).
Hemi-epiphytic species can be hemi-epilithic on exposed rocky surfaces.
Some species of Ficus are more or less clearly rheophytic (see FM17-2: 27), e.g.,
F. ischnopoda, F. macrostyla, and F. squamosa, the latter two with fruitlets with long
persistent styles and retrorse stiff hairs on the style and the margin of the fruit body
morphologically adapted to the unusual substrate.
In general, the upper altitudinal limit of lowland species is 1500 m. Montane species,
above 2000 m, are found in Ficus and Streblus. Morus macroura can also be regarded
as montane. The hardier Moraceae of Sino-Himalaya do not reach Malesia.
REPRODUCTIVE BIOLOGY
Pollination — Wind-pollination occurs in Moraceae with urticaceous type stamens and

may occur in some other species, such as those with long pendulous staminate inflores-
cences. Anemophily is also reported for patent (scentless) staminate inflorescences of
some species of Artocarpus (e.g., A. elasticus and A. rigidus (Corner 1940, 1988; Jarrett
1959) that give off clouds of pollen. The ballistic release of pollen from urticaceous
type stamens allows plants to inhabit the forest undergrowth where they can make use
of weak air currents, as above streaming water, for the transport of pollen. This type
of pollination is described and discussed for Streblus pendulinus and Trophis scandens
by Williams & Adam (1993).
Geitonogamy might occur in species with bisexual inflorescences, such as Dorste
nia.
Insect-pollination is probably the predominant mode of pollination in the family. The
unique mode in Ficus is well-documented (see FM 17-2: 51– 54). Another is described
for Artocarpus (Van der Pijl 1953), in which species with staminate inflorescences (e.g.,
A. dadah and A. integer; Corner 1940) emit a sweet scent of honey and burnt sugar, to
attract small flies and beetles which subsequently breed in the inflorescence. Pollina-
tion based on insects breeding in staminate inflorescences is common in many tropi-
cal plant groups with reduced and unisexual flowers (such as palms), and is probably
widespread in those Moraceae with dense unisexual inflorescences as well. In Castilla
elastica and Antiaropsis decipiens this role is played by thrips (Sakai 2001; Zerega et
al. 2004), which could be the case in other genera where the staminate inflorescences
are (±) closed before anthesis. In some species of Artocarpus with scentless staminate
inflorescences, clouds of pollen are given off, e.g. in A. elasticus and A. rigidus (Corner
1938, 1940; Jarrett 1959).
An unusual mode of pollination is described for A. integer in Sarawak (Sakai et al.
2000): staminate inflorescences infected and covered by fungus are visited by a spe-
cies of gall midge that feeds on the mycelium and oviposits on the inflorescence. The
midges transport pollen to pistillate inflorescences. It is not clear whether this is a local
phenomenon or widespread in the species.
Dispersal — The seeds are usually animal dispersed, although this may not be the case
if the fruits are dehiscent drupes. Here, small endocarp bodies (kernels) are ejected
(Dorstenia and Fatoua) or, if large, are squeezed out (Antiaropsis and Streblus p.p.),
and drop on the forest floor or into water (by which they are carried on) or are ejected
(see p. 8). The endocarp bodies, which are animal dispersed, remain inside the infructes
cence (Broussonetia and Ficus). Alternatively they may be large, black, and embedded
in structures with contrasting colours (e.g. white of the exocarp and red of tepals as
in Antiaropsis), in which case they may be dispersed by birds. In the majority of the
Moraceae the fruits (and seeds ) are enclosed in fleshy structures consisting of connate
perianths and interfloral bracts (Artocarpus and related genera), or also adnate to the
receptacle (Hullettia), or enclosed in fleshy receptacles (Ficus). The infructescence may
more simply be just aggregations of flowers in fruit of which the fleshy perianths are
the attractive parts (Morus). More or less solitary fleshy (dehiscent or indehiscent) fruits
occur in Streblus. The colour of individual fruits or of infructescences varies from red
to orange. Large infructescences are often greenish to yellowish. The infructescences of
Ficus can be blackish, or sometimes brown or purplish and then taken by fruit bats.
Moraceous fruits and infructescences are taken by birds, monkeys, squirrels, and
other arboreal animals by day. By night the same fruits, or those of allied species, are
taken by nocturnal animals, such as fruit bats and civet cats. Cauliflorous species of
Artocarpus and Ficus may also be dispersed by bats as well as by ground mammals,
ranging from elephants and rhinoceros to pigs and mouse deer.
Water may play a role in distribution of diaspores ejected or fallen into streams. The
fruitlets of some rheophytic Ficus species are adapted to attachment to the substrate.
References: Corner, E.J.H., Notes on the systematy and distribution of Malayan phanerogams, II.
The jack and the chempedak. Gard. Bull. Singapore 10 (1938) 56 – 81. — Corner, E.J.H., Wayside
trees of Malaya ed. 1, 1. Government Printing Office, Singapore (1940). — Corner, E.J.H., Wayside
trees of Malaya ed. 3, 1. Malayan Nature Society, Kuala Lumpur (1988). — Jarrett, F.M., Studies in
Artocarpus and allied genera, III. A revision of Artocarpus subgenus Artocarpus. J. Arnold Arbor. 40
(1959) 113 –115. — Sakai, S., Thrips pollination of androdioecious Castilla elastica (Moraceae) in a
seasonal tropical forest. Amer. J. Bot. 88 (2001) 1527–1534. — Sakai, S., M. Kato & H. Nagamasu,
Artocarpus (Moraceae) gall midge pollination mutualism mediated by a male-flower parasitic fungus.
Amer. J. Bot. 87 (2000) 440 – 445. — Van der Pijl, L., On the flower biology of some plants from Java
– with general remarks on fly-traps (species of Annona, Artocarpus, Typhonium, Gnetum, Arisaema
and Abroma). Ann. Bogor. 1 (1953) 77– 99. — Williams, G. & P. Adam, Ballistic pollen release in
Australian members of the Moraceae. Biotropica 25 (1993) 478 – 480. — Zerega, N.J.C., L.A. Mound
& G.D. Weiblen, Pollination in the New Guinean endemic Antiaropsis decipiens is mediated by a new
species of thrips, Thrips antiaropsidis spec. nov. (Thysanthoptera: Thripsidae). Int. J. Pl. Sci. 165 (2004)
1017–1026.
DELIMITATION AND SUBDIVISION OF THE FAMILY
In the classification by Engler (1888) the family Moraceae included the subfamily
Conocephaloideae, with six genera, including the Asian genus Conocephalus Blume.
In a revisional study by Chew (1963) the genus was united with the urticaceous genus
Poikilospermum Zipp. ex Miq. This led to proposals to transfer some or all the genera
of the Conocephaloideae (Chew 1963 and Corner 1962, respectively) to the Urticaceae.
For these six genera the family Cecropiaceae Berg (1978) was established.
The order Urticales constitute a clear-cut and coherent group. Moraceae differs
from the Urticaceae in the presence of milky sap (or latex), the apical attachment of
the ovule, the common presence of two stigmas, the absence of elongate cystoliths, and
the predominantly woody habit in Urticaceae. The stamens are always inflexed in the
bud and bend outwards suddenly, throwing the pollen into the air. Urticaceous stamens
also occur in most species of the tribe Moreae. Inflexed stamens resembling those of
Urticaceae occur in some genera of the Ulmaceae. The family Cecropiaceae is entirely
woody and differs from the Moraceae in some features shared with Urticaceae, such as
the basally attached ovule and single stigma, by strict dioecism, the absence of milky
sap, the presence of adventitious roots (in the woody genera of Moraceae only present
in Ficus), and the always spirally arranged leaves. Ulmaceae share with Moraceae the
apically attached ovule, but lack milky sap. In contrast to the other families, the flowers
are morphologically and often also functionally bisexual. Judd et al. (1994) suggested
including Cannabidaceae, Cecropiaceae, Moraceae, in the Urticaceae, leaving the Celti-
daceae and Ulmaceae as two much smaller urticalean families.
The Urticales show rather clear morphological affinities to the Malvales and some-
what remotely to the Euphorbiales, but they do not show links to the other families
traditionally ranked among the Hamamelidae. Molecular studies place Urticales among
the Rosales (Sytsma et al. 2002), but this contradicts patterns of morphological differ-
entiation, ecology, and phytogeography, which suggest a different evolutionary history
of the group (but see chapter ‘Phytochemistry and Chemotaxonomy’, p. 11).
For the reduced family Moraceae Corner (1962) proposed seven tribes, which with
some adjustments was reduced to five tribes (Berg 1988, 2001): Moreae (8 genera,
70 –75 spp., centred in Asia and characterized by the urticaceous type of stamen, see
p. 7); Castilleae (8 genera, 50 – 60 spp., centred in the Neotropics and characterized by
trees with the architectural model of Cook (see p. 138); Dorstenieae (8 genera, 125 –130
spp., amphi-atlantic and characterized by circular bisexual inflorescences); and Ficeae
(1 genus, 720 –750 spp., characterized by essentially bisexual, urceolate inflorescences).
More recently the tribe Artocarpeae was redefined and two additional tribes established
(Berg 2005): Artocarpeae (4 genera, c. 55 spp, centred in Asia and characterized by
many-seeded infructescences, mostly formed by connate flowers, but with free fruits);
Antiaropsideae (2 genera, 3 or 4 spp.), in New Caledonia and New Guinea and charac-
terized by involucrate inflorescences and dehiscent drupes); and Soroceae (5 genera, 23
spp., neotropical and characterized by simply constructed, Moreae-like inflorescences

but with stamens straight in the bud).
In the classification of Moraceae proposed by Corner (1962) the number of genera
previously recognised for the Asian–Australasian region was reduced considerably.
This study and others by Berg (e.g., 1986, 1988), in which some of the Asian–Aus-
tralasian representatives of the family were involved, resulted in the recognition of 37
genera worldwide. These appear to be largely natural ones and show interesting patterns
in geographical distribution and taxonomic relationships.
A molecular phylogenetic study by Datwyler & Weiblen (2004) largely supports
distinctness and homogeneity of Castilleae, Dorstenieae, and Ficeae, but not so of Arto-
carpeae (sensu stricto), Moreae, and Soroceae, not even at the generic level. Moreover,
the study indicates affinities of Antiaropsideae to Castilleae and Ficeae. Support for
this phylogeny from morphology and phytogeography is still wanting. The phylogeny
of the more recent molecular study by Zerega et al. (2005) is rather similar and pays
attention to biogeography and divergence times, but hardly to patterns in morphological
differentiation.
References: Berg, C.C., Cecropiaceae a new family of the Urticales. Taxon 27 (1978) 39 – 44.
— Berg, C.C., The delimitation and subdivision of the genus Maclura (Moraceae). Proc. Kon. Ned.
Akad. Wetensch. C, 89 (1986) 241– 247. — Berg, C.C., The genera Trophis and Streblus (Moraceae)
remodelled. Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 345 – 362. — Berg, C.C., Flora Neotropica
Monograph 83. Moreae, Artocarpeae, Dorstenia (Moraceae). With introductions to the family and Ficus
and with additions and corrections to Flora Neotropica Monograph 7. New York (2001) — Berg, C.C.,
Flora Malesiana precursor for the treatment of Moraceae 8: other genera than Ficus. Blumea 50 (2005)
535 – 550. — Chew, W.-L., Flora malasianae precursores XXXIV. A revision of the genus Poikilo
spermum (Urticaceae). Gard. Bull. Singapore 20 (1963) 1–103. — Corner, E.J.H., The classification
of Moraceae. Gard. Bull. Singapore 19 (1962) 187–252. — Datwyler, S.L. & G.D. Weiblen, On the
origin of the fig: phylogenetic relationships of Moraceae from ndhF sequences. Amer. J. Bot. 91 (2004)
767–777. — Engler, G.H.A., Moraceae, in: G.H.A. Engler & K. Prantl, Natürliche Pflanzenfamilien
3, 1 (1888) 66 – 98. Engelmann, Leipzig. — Judd, W.S., R.W. Sanders & M.J. Donoghue, Angiosperm
family pairs: preliminary phylogenetic analyses. Harvard Pap. Bot. 5 (1994) 1– 51. — Sytsma, K.J.,
J. Morawetz, J.C. Pires, M. Nepokroeff, E. Contoi, M. Zjhra, J.C. Hall & M.W. Chase, Urticalean
rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences.
Amer. J. Bot. 89 (2002) 1531–1546. — Zerega, N.J.C., W.L. Clement, S.L. Datwyler & G.D. Weiblen,
Biogeography and divergence times in the mulberry family (Moraceae). Molec. Phylogenet. Evol. 37
(2005) 402 – 416.
USES
Streblus elongatus and several species of Artocarpus are, or have been, outstanding
producers of timber. Lesser known are the timber trees found in the genera Antiaris,
Parartocarpus, and Prainea (Boer & Sosef 1998). Ficus elastica and the American
Castilla elastica have been used for rubber and remains of their plantations may be
found throughout Malesia, but they have given place to Hevea because of their resin-
ous milksap. Poisonous milksap occurs in the famous upas tree Antiaris (see Boer et
al. 1999) and in the much less common Parartocarpus; that of both is used as (one of
the components of) dart and arrow poison in South-east Asia. The sap of many species
of Artocarpus, by contrast, being very sticky and innocuous, is used for birdlime and
general adhesive; whereas that of A. lowii, is oily and greasy and is used as an ointment
and for cooking. Fibrous and easily-stripped bark of several species of Artocarpus,
such as A. elasticus, is made into coarse bark cloth and binding material, but this is dis
appearing even from the life of jungle folk; many anthropological exhibits in museums
are held together by this material. The fibrous bark of Ficus is used for string, even for
bow strings, but being readily cropped these plants have not been commercialised. Paper
is manufactured from the inner bark of the paper mulberry Broussonetia papyrifera,
as well as fibre clothing and string among primitive people (Berg 2003). Edible fruits
are the mulberries (Morus), various kinds of figs (Ficus), several kinds of Artocarpus,
and indeed Antiaris, the fruits of which seem always devoid of poison. Introduced
mulberries have found little favour in Malesia. The cultivated fig (Ficus carica) is
rarely seen because it succumbs readily to insect attacks. Sometimes the Indochinese
Ficus auriculata is cultivated, but its fruit is inferior. Certain wild figs are, however, not
despised. The best known fruit trees of the family in Malesia are the jackfruit (Artocar
pus heterophyllus) and the chempedak or chemedak (A. integer). They are grown both
for the edible pulp around the seeds and for the seeds themselves which are roasted.
Some varieties of the jack produce infructescences, which may hold the world record
for fruit size. Few fruits can equal in stench some varieties of the chempedak. The
breadfruit tree (Artocarpus altilis), commonly seen in villages, is extremely abundant
both wild and cultivated in New Guinea, where it is grown for the unripe fruit which is
baked, roasted, or boiled more like a tuber than a fruit; the seedless form is most usually
cultivated. Artocarpus odoratissimus is cultivated in Borneo and the Philippines for
the sweet pulp round the seed. Artocarpus nitidus, with the uniformly succulent kind
of infructescence, is sometimes found in gardens in West Malesia, where it is used also
for jam and conserves. Other species of Artocarpus are mostly wild trees which have
been spared from felling for sake of their fruit. Leaves and young figs of several Ficus
species are eaten either cooked or raw as vegetables.
Yellow dye is extracted from wood of Maclura cochinchinensis (Heyne 1927).
References: Berg, C.C., Broussonetia papyrifera (L.) L’Hér. ex Vent., in: M. Brink & R.P. Escobin
(eds), Plant Resources of South-East Asia 17, Fibre plants (2003) 91– 95. — Boer, E., M. Brink &
M.S.M. Sosef, Antiaris toxicaria Lesch., in: L.S. de Padua, N. Bunyapraphatsara & R.H.M.J. Lemmens
(eds), Medicinal and poisonous plants 1. Plant Resources of South-East Asia 12 (1) (1999) 126 –129.
Backhuys Publishers, Leiden. — Boer, E. & M.S.M. Sosef, Antiaris, Parartocarpus, and Prainea, in:
M.S.M. Sosef, L.T. Hong & S. Prawirohatmodjo (eds), Timber trees: Lesser known timbers. Plant
Resources of South-East Asia 5 (3) (1998) 73 –75, 425 – 427, 469 – 470. Backhuys Publishers, Leiden.—
Heyne, K., De nuttige planten van Nederlandsch-Indië ed. 2, 1 (1927). Ruygrok & Co., Batavia.
KEY TO THE TRIBES
1a. Inflorescences urceolate, the opening entirely closed by (ostiolar) bracts; waxy
glandular spots on the lamina lower surface (on the base of the midrib, in the axils
of lateral veins and/or furcations of lateral veins), or on leafy twigs just below the
nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ficeae
b. Inflorescences mostly spicate, capitate, or discoid, or if urceolate, then the open-
ing not entirely closed; waxy glandular spots mostly absent (present in Artocarpus
subg. Pseudojaca) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Stamens bent inwards in bud, at anthesis bending outwards suddenly and elasti-
cally, or if straight in the bud, then the plant armed with thorns on the branches . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moreae
b. Stamens straight or bent outwards in bud or, if bent inwards, then straightening
gradually; plants without thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Inflorescences discoid-, turbinate- or globose-capitate, bisexual, or if (secondarily)
unisexual, then fruits adnate to the perianth (and receptacle), or the plant herba-
ceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dorstenieae
b. Inflorescences spicate, clavate- to globose- (to discoid-)capitate, or discoid-capitate
to urceolate and involucrate, (primarily) unisexual; plant woody . . . . . . . . . . . . . 4
4a. Trees with self-pruning branches; inflorescences discoid and involucrate with im-
bricate bracts; fruit adnate to the perianth, if not so, then the exocarp indehiscent
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castilleae
b. Trees without self-pruning branches; inflorescences not involucrate (or subinvolu-
crate), or if involucrate, with imbricate bracts and the fruits with a dehiscent exo-
carp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Inflorescences discoid to urceolate, involucrate with basally attached imbricate
bracts enclosing the flowers, at least before anthesis; fruits dehiscent drupes . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antiaropsideae
b. Inflorescences spicate to clavate-, ellipsoid-, to globose-, or discoid-capitate, if
the heads subtended by bracts, then these short and not enclosing flowers before
anthesis; fruits indehiscent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Artocarpeae
KEY TO THE GENERA
1a. Plants herbaceous or suffrutescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b. Plants woody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Plants branched; sap watery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fatoua
b. Plants monocaul and subacaul; sap milky . . . . . . . . . . . . . . . . . . . . . . . Dorstenia
3a. Plants armed with thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
b. Plants unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4a. Staminate and pistillate inflorescences globose-capitate; stamens straight in the
bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maclura
b. Staminate inflorescences spicate, racemose or capitate and stamens inflexed in the
bud; pistillate inflorescences with 1– 3 flowers . . . . . . . . . . . . . . . . . Streblus p.p.
5a. Inflorescences urceolate and stigmas and anthers not exposed at anthesis; waxy
glandular spots on the lamina lower surface (at the base of the midrib, in the axils
of lateral veins and/or furcations of lateral veins), or on the nodes of leafy twigs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ficus
b. Inflorescences not urceolate and stigmas and anthers exposed at anthesis; waxy
glandular spots mostly lacking, but present on the lamina beneath in the axils of
lateral veins only in Artocarpus subg. Pseudojaca . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Stipules fully amplexicaul . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Stipules not fully amplexicaul to lateral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7a. Stipules connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castilla

b. Stipules free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Artocarpus p.p.
8a. Leaves of lateral (flower-bearing) branches spirally arranged (or subopposite)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
b. Leaves of lateral (flower-bearing) branches distichous . . . . . . . . . . . . . . . . . . . 13
9a. Stipules connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parartocarpus
b. Stipules free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Pistillate flowers connate; staminate flowers with 1 stamen . . . . Artocarpus p.p.
b. Pistillate flowers free; staminate flowers with 2 – 4 stamens . . . . . . . . . . . . . . . 11
11a. Inflorescences discoid- to turbinate-capitate with marginal bracts . . . . Hullettia
b. Inflorescences spicate or globose-capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Lamina with 3 or 4 pairs of lateral veins (S. ascendens) . . . . . . . . . . . . . Streblus
b. Lamina with 5 – 9 pairs of lateral veins (B. papyrifera) . . . . . . . . . Broussonetia
13a. Inflorescences bisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
b. Inflorescences unisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14a. Inflorescences cymose; perianth of the pistillate flower tubular . . . . Bleekrodea
b. Inflorescences spicate; perianth of the pistillate flower with free imbricate
tepals (S. elongatus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streblus
15a. Inflorescences discoid-capitate or uniflorous, involucrate with basally attached
imbricate bracts, or bracts concentrated in the upper part of the receptacle of the
pistillate inflorescence with a single flower adnate to the receptacle . . . . . . . . 16
b. Inflorescences globose-capitate, spicate, racemose, cymose, or if uniflorous, then
not involucrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16a. Receptacle of the pistillate inflorescence infundibuliform, enclosing and fused
with a single flower; involucral bracts small and partly scattered on the receptacle;
staminate inflorescences usually several together . . . . . . . . . . . . . . . . . . Antiaris
b. Receptacle of the pistillate inflorescence discoid, the flower(s) free and the invo
lucral bracts large, in 4 or 5 rows imbricate; staminate inflorescences solitary . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antiaropsis
17a. Scaled axillary resting buds present; apices of branches abscised, elongation of
the branch by the terminal apical lateral bud . . . . . . . . . . . . . . . . . . . . . . . . . . 18
b. Scaled resting buds absent (or weakly developed) and apical bud persistent . . 19
18a. Pistillate inflorescences globose-capitate; stigma 1; tepals of the staminate flower
valvate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Broussonetia
b. Pistillate inflorescences spicate to subcapitate; stigmas 2; tepals of the staminate
flower imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morus
19a. Staminate material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
b. Pistillate material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
20a. Stamen 1, straight in the bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
b. Stamens 3, 4, or 5, inflexed in the bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
21a. Stipules lateral; heads globose to obovoid; perianth 2-lobed to subentire . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prainea
b. Stipules fully amplexicaul or, if lateral, then heads usually distinctly longer than
wide and perianth 2– 4-lobed to parted . . . . . . . . . . . . . . . . . . . . . . . Artocarpus
22a. Inflorescences spicate, all flowers sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

b. Inflorescences racemose or cymose or subcapitate, all or some of the flowers
pedicellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
23a. Tepals valvate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
b. Tepals imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24a. Midrib of the lamina impressed above; stamens 4 . . . . . . . . . Broussonetia p.p.
b. Midrib of the lamina prominent above; stamens 3 (or 4) (T. scandens) . . Trophis
25a. Bracts basally attached . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streblus p.p.
b. Bracts (partly) peltate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26a. Midrib prominent above (T. philippinensis) . . . . . . . . . . . . . . . . . . . . . . Trophis
b. Midrib impressed above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streblus p.p.
27a. Inflorescences cymosely branched . . . . . . . . . . . . . . . . . . . . . . . . . . Bleekrodea
b. Inflorescences unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streblus p.p.
28a. Flowers connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Artocarpus
b. Flowers free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29a. Tepals free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streblus
b. Tepals connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Stigma 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Broussonetia
b. Stigmas 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31a. Perianth fleshy in fruit; ovary adnate to the perianth, or if not, then the plant
scandent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trophis
b. Perianth membranaceous in fruit; fruit free and the plant a shrub . . Bleekrodea
Tribe Moreae
Moreae Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509; Corner, Gard. Bull. Singapore 19 (1962)
211; C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 345.
Broussonetieae Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 508.
Strebleae Bureau in DC., Prodr. 17 (1873) 215.
Fatoueae Engl., Nat. Pflanzenfam. 3, 1 (1888) 71.
Trees, shrubs, or in Fatoua, herbs or subshrubs, dioecious or sometimes monoecious,

with or without uncinate hairs. Leaves alternate and distichous, or less commonly
spirally arranged, sometimes (sub)opposite; stipules small to large, lateral to semi-
amplexicaul. Inflorescences unisexual or less commonly bisexual, racemose, spicate,
cymose, capitate, or uniflorous pedunculate, bracteate; staminate flowers with 3, 4, or
5 tepals; stamens 3, 4, or 5, inflexed in the bud and bending outwards suddenly and
elastically at anthesis, or sometimes straight in the bud; pistillode usually present;
pistillate flowers with 4, free or connate, tepals; ovary mostly free, stigmas 2 and then
equally or unequally long, or 1. Fruit drupaceous, dehiscent and releasing or ejecting
the endocarp body, or indehiscent, achene-like, or forming a drupaceous whole with
the fleshy perianth. Seed small and with endosperm, or large and endosperm (almost)
lacking; testa mostly with a thickened vascularised part below the hilum; embryo vari-
ous.
Distribution — This tribe comprises eight genera, seven of them represented in
Malesia. The eighth genus is Milicia Sim, with two species, is confined to the African
continent. The number of species is c. 70, c. 47 of them occur in the Asian–Australasian

region. Nine species occur in the Neotropics and two are confined to the North Ameri-
can continent. Twelve species occur in the African region, seven of them are endemic
to Madagascar and adjacent islands. These numbers show that the Asian–Australasian
region is the centre of this tribe. The largest genus in this region is Streblus.
The genera Morus and Broussonetia can be regarded as elements of the northern
warm temperate region (see below). Some species of Maclura and Streblus, most of
them thorn-bearing shrubs, small trees or climbers, are elements of relatively dry cli-
matic conditions.
Staminate flowers and pollination — This tribe is characterised by stamens similar

to those characteristic of Urticaceae, but are absent in Maclura sect. Cudrania. The
stamens are inflexed in the flower bud, and bend outwards suddenly and elastically,
throwing pollen in the air. This mechanism of ballistic release of pollen requires a
well-developed perianth and sufficient space in the inflorescence for the opening of the
perianth. In contrast to other tribes, pistillodes are commonly present. They are often
quadrangular and apparently play a role in keeping the stamens under tension until
anthesis. In the Moreae the inflorescences are relatively simply constructed, and the
staminate flowers at least not densely packed – a connection with the ballistic release
of pollen. The perianth and the number of stamens is not reduced as often happens in
other tribes and the relatively simple construction of the pistillate inflorescences could
be related to the requirement of simple staminate inflorescences. Pistillate flowers are
mostly free, but (partly) connate in some subdivisions of Maclura: sect. Plecospermum
and sect. Cudrania ser. Connatae.
Habit — The habit varies from tall primary forest trees to small trees and shrubs, sub-
shrubs, to herbs (in Fatoua), and climbers (in species of Maclura and Trophis scandens).
Thorns are found in the species of Maclura and in some species of Streblus, in the
former genus they facilitate climbing. Milky sap is absent in Fatoua.
Inflorescences — The inflorescences of the Moreae are mostly unisexual. Bisexual

inflorescences are typical for Bleekrodea, Fatoua, and Streblus elongatus, although in
these taxa the inflorescences can be unisexual, pistillate and/or staminate. In the other
taxa the inflorescences are normally unisexual and the plants mostly dioecious. The
inflorescences are short-branched cymes, racemes, spikes, heads, or uniflorous (as the
pistillate ones in several species of Streblus).
Fruits — Dehiscent drupes occur in Bleekrodea, Broussonetia, Fatoua, and some of

the Streblus species. In other Streblus species the fruits are red or black coloured ber-
ries. In the other genera the fruits are largely or entirely enveloped by fleshy, mostly
coloured perianth.
Delimitation — The tribe as currently defined includes most of the genera referred
to this tribe by Corner (1962), but it also includes the other genera with urtica-
ceous stamens such as Broussonetia, Maclura, and Malaisia, as well as the neotropical
genus Olmedia. The latter two have been reduced to sections of Trophis (Berg 1988).
However, the neotropical genera Clarisia and Sorocea are currently included in the
tribe Soroceae (see Berg 2005).
References: Berg, C.C., The genera Trophis and Streblus (Moraceae) remodelled. Proc. Kon. Ned.
Akad. Wetensch. C, 91 (1988) 345 – 362. — Berg, C.C., Flora Malesiana precursor for the treatment of
Moraceae 8: other genera than Ficus. Blumea 50 (2005) 535 – 550. — Corner, E.J.H., The classification
of Moraceae. Gard. Bull. Singapore 19 (1962) 187– 252.
KEY TO THE GENERA OF THE MOREAE
1a. Plants herbaceous (or suffrutescent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fatoua

b. Plants woody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Shoot apices deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
b. Shoot apices persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3a. Tepals of the pistillate flowers free; stigmas 2, equal . . . . . . . . . . . . . . . . Morus
b. Tepals of the pistillate flowers connate; stigma 1, or if 2, then unequal . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Broussonetia
4a. Plants armed with curved or straight spines; pistillate inflorescences globose-capi-
tate; tepals and bracts mostly with yellow dye accumulated in immersed glands . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maclura
b. Plants unarmed, or if armed, then with straights spines; the pistillate inflorescences
not globose-capitate and tepals and bracts without yellow dye containing glands
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Plants mostly with thorns or lamina scabrous above; inflorescences spicate or uni-
sexual; tepals of the pistillate flowers (almost) free . . . . . . . . . . . . . . . . . Streblus
b. Plants without thorns, lamina smooth above; inflorescences cymose if bisexual;
tepals of the pistillate flowers connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Inflorescences bisexual, cymose; tepals of the staminate flowers 4 or 5 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bleekrodea
b. Inflorescences unisexual, spicate; tepals of the staminate flowers 4 or 3 . Trophis
MORUS
Morus L., Sp. Pl. (1753) 986; Endl., Gen. Pl. (1837) 278; Gen. Pl., Suppl. 4 (1848) 33; Bureau in DC.,
Prodr. 17 (1873) 237; Baill., Hist. Pl. 6 (1875) 190; Benth. & Hook.f., Gen. Pl. 3 (1880) 364; Engl.
in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 72; Nakai, J. Arnold Arbor. 8 (1927) 234; Gagnep.,
Fl. Indo-Chine 5 (1928) 706; C.C. Berg, Fl. Neotrop. Monogr. 83 (2001) 25.
Morophorum Neck., Elem. Bot. 3 (1790) 255.
Trees, dioecious, unarmed, shoot apices shed, with axillary scaled resting buds.
Leaves distichous; lamina pinnately veined or trinerved to triplinerved, with cysto-
liths usually only above; stipules free, almost lateral. Inflorescences unisexual, usually
solitary on the lower leafless nodes of new (short-)shoots arising from well-developed
scaly resting buds on wood of the previous season, ebracteate or bracteate. Staminate
inflorescences spicate (or racemose); perianth 4-parted, segments imbricate; stamens
4, inflexed in the bud. Pistillate inflorescences capitate to spicate; perianth 4-parted;
ovary free, style subterminal, stigmas 2. Fruiting perianth enlarged, more or less suc-
culent; fruit with a broad base, exocarp thin-fleshy, thicker on the seed-bearing side,
indehiscent, endocarp crustaceous with a woody plug towards the hilum; seed with
endosperm, embryo curved, cotyledons equal and flat, not enclosing the long radicle.
Distribution — The genus comprises ten to fifteen species, most of them in temper-
ate to subtropical regions of the northern hemisphere, some in montane habitats in the
tropics (M. insignis Bureau in the Neotropics and M. macroura in SE Asia and Malesia),
one in the tropical lowlands of Africa (M. mezozygia A. Chev., as evergreen tree in rain-
forest or as deciduous tree in drier conditions), and one in dry regions of SE United
States and Mexico (M. celtidifolia Kunth). The North American M. rubra L. is closely
related to the group of species associated with northern temperate to subtropical condi-
tions and centred in the Himalayan and East Asian region and constituting a complex
of very closely related taxa for which a recent taxonomic evaluation is still wanting;
this complex comprises, e.g., M. alba L. and M. nigra L.
Morphology — The association with northern temperate conditions is morphologi-
cally expressed in the abscission of shoot apices, the presence of ± well-developed
scaled resting buds, and deciduousness, features which are at least partly still pertained
by the tropical lowland species.
Chromosomes — Polyploids (4x, 6x, 8x, or even 22x) are found in the Himalayan-
East Asian complex of taxa.
Subdivision — Leroy (Bull. Mus. Hist. Nat. (Paris), Sér. 2, 21: 725) recognised three
subgenera: Afromorus (with M. mesozygia), Gomphomorous (with M. insignis), and
Eumorus with the rest of the species.
Uses — The genus has a very long historical association with mankind for edible
fruits of many species and for the cultivation of the slik-moth.
KEY TO THE SPECIES
1a. Lower surface of the lamina barbate, with tufts of hairs in the axils of lateral veins
and major furcations of lateral veins; stipules 0.4 –1.2 cm long; pistillate spikes to
3.5 cm long, in fruit black or blackish red. — Widespread, in cultivation . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. M. alba
b. Lower surface of the lamina not barbate, hairs ± evenly distributed on the veins;
stipules 1– 2.5 cm long; pistillate spike (1–)4 – 8 cm long, in fruit greenish or yel-
lowish. — Sumatra, Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. M. macroura
1. Morus alba L.
Morus alba L., Sp. Pl. (1753) 986; Ser., Descr. Cult. Muriers (1855) 19, t. 1–18; Bureau in DC., Prodr.
17 (1873) 238.
Tree up to 10(–15) m tall. Leafy twigs 1– 4 mm thick, (minutely and/or sparsely)

puberulous; resting buds 0.3 – 0.8 cm long. Leaves distichous; lamina ovate or subovate
(in outline), 10 – 20 by 6 –15 cm, entire or 3 – 5(– 9)-lobed to -fid, apex acuminate, base
rounded to cordate, margin crenate- to serrate-dentate; upper surface minutely white
puberulous to hispidulous, scabridulous to smooth, lower surface puberulous to hispidu-
lous on the (main) veins, with concentrations of hairs in the axils of lateral veins and
furcations of lateral veins, scabridulous to smooth; lateral vein (3 –)4 – 6(–7) pairs, the
basal pair up to 1/2(– 2/3) the length of the lamina, branched, the other lateral veins
also branched or furcate far from the margin, tertiary venation loosely scalariform;
petiole 1– 6 cm long, 2 – 2.5 mm thick, minutely puberulous; stipules 0.5 –1.2 cm long,
puberulous, caducous (or subpersistent). Staminate inflorescences axillary (often in the
axils of scale leaves at the base of new shoots), solitary or in pairs, spicate to racemose,
ebracteate; peduncle 0.5 –1.5 cm long, minutely white puberulous to tomentellous; spike
0.5 – 2 cm long; flowers 5 – 20, sessile or up to 1.5 mm long pedicellate; perianth 1– 2
mm long, minutely white puberulous, stamens 2 – 3.5 mm long, anthers c. 0.5 mm long.
Pistillate inflorescences axillary (often in the axils of scale leaves at the base of new
shoots), solitary (or in pairs), spicate to subcapitate, ebracteate; peduncle 0.5 –1.5 cm
long, white puberulous to tomentellous; spike 0.5 – 2 cm long; flowers 15 – 50; tepals
1–1.5 mm long, sparsely white puberulous or only ciliolate; ovary c. 1 mm long, style
0 –1.5 mm long, stigmas 1.5 – 3.5 mm long. Infructescences 0.8 – 3.5 cm long, 0.5 –1.5
cm thick, reddish to black.
Distribution — From Afghanistan through the Himalaya region to China and Japan,
wild or cultivated; in Malesia: introduced and cultivated in Sumatra (also Simalur),
Malay Peninsula, Java, Borneo, Philippines, Celebes, Lesser Sunda Islands (Lombok,
Timor), Moluccas (Ambon, Tanimbar Islands), New Guinea; naturalised in parts of Java
and N Luzon.
Habitat — Grown at altitudes up to 2000 m.
Uses — This species is in cultivation for its fruits.
Notes — Numerous names at the rank of species and below that rank have been
attached to material of the genus Morus with minor morphological differences. This
was a reason for Bureau (1873) to include them in M. alba. This broadly construed
species also comprised material (with long pistillate inflorescences) under the name
var. laevigata that is now included in F. macroura. For the remainder (with short pis-
tillate inflorescences) two groups of varieties and subvarieties were recognized: those
with short (0 – c. 0.5 mm long) and those with long (c. 0.5 –1.5 cm) styles. In material
introduced outside the natural range to various parts of the tropics, these two categories
can also be recognized, the former in general in plantations for silk production and
temperate climatical conditions, the latter especially in trees used as fruit trees and in
tropical lowland. A good number of specimens with pistillate inflorescences cannot be
included in one of the two forms because of ± intermediate style lengths. Due to the
lack of correlated characters in staminate inflorescences and in vegetative parts the best
provisional solution could be to treat the form with short styles as var. alba and the
other as var. indica (L.) Bureau (in DC., Prodr. 17 (1873) 243, based on Morus indica
L. (1753) 986). However, a thorough study on the Morus alba complex with traditional
and modern methodology is needed to arrive at its taxonomic understanding. For this
reason no synonyms are listed.
The long-styled material is often identified as Morus australis Poir., according to
Bureau (1873) based on material of Broussonetia papyrifera (L.) Vent. The name has
been reintroduced for Morus material by Rehder (J. Arnold Arbor. 10 (1929) 123) and
since often applied for long-styled material in cultivation mainly in lowland tropics.
2. Morus macroura Miq.

Morus macroura Miq., Pl. Jungh. (1851) 42; Fl. Ned. Ind. 1, 2 (1859) 280; Bureau in A.DC., Prodr.
17 (1873) 247; Boerl., Bijdr. Fl. Sum. (1884) 32; Koord., Exkurs.-Fl. Java 2 (1912) 84, f. 4F; Atlas
Baumart. Java (1924) t. 745; Corner, Gard. Bull. Singapore 19 (1962) 215; Backer & Bakh.f., Fl.
Java 2 (1965) 15. — Morus alba L. var. laevigata Wall. ex Bureau in A.DC., Prodr. 17 (1873) 245.
— Morus laevigata (Wall. ex Bureau) Hook.f., Fl. Brit. India 5 (1888) 492; Gagnep., Fl. Indo-Chine
5 (1928) 708; Bor, Man. Ind. For. Bot. (1953) 137.
Tree up to 32 m tall, deciduous. Leafy twigs 1.5 – 3 mm thick, puberulous to hirtel-

lous to subtomentose; resting buds 0.3 –1 cm long. Leaves distichous; lamina ovate to
subovate to elliptic, 6 –16(– 23) by 4 –11(–14) cm, entire (or when juvenile 3-lobate),
chartaceous to subcoriaceous, apex acuminate to acute, base subcordate to rounded (to
subcuneate), margin serrate-denticulate (to subentire); upper surface hispidulous to
puberulous, mainly on the main veins, ± scabrous to smooth; lower surface puberulous
to subtomentellous on the veins; lateral veins 3 – 6 pairs, tertiary venation scalariform;
petiole 1.5 – 6 cm long, puberulous to hirtellous; stipules 1–1.5 cm long, sparsely to
e 1 mm
3 cm
Fig. 1. Morus macroura Miq. a. Leafy twig with axil-

lary resting bud; b. staminate flower with pistillode;
b a c. pistillate flower; d. fruit; e. endocarp bodies (a: Raub 1;
2 mm b, c, e: Bosscha 2; d: Plasschaert s.n., Sumatra).
Map 1. Distribution of Morus macroura Miq.
densely puberulous to hirtellous, caducous (or subpersistent). Staminate inflorescences

axillary (often in the axils of scale leaves at the base of new shoots), solitary or in pairs,
spicate (or racemose); peduncle 0.4 –1.5 cm long, puberulous; spike (1–)4 – 8 cm long;
flowers numerous; pedicel 0 –1.5 mm long; perianth c. 1.5 mm long, puberulous to
tomentellous; stamens c. 2.5 mm long, anthers c. 0.8 mm long. Pistillate inflorescences
axillary, (often in the axils of scale leaves at the base of new shoots), solitary (or in
pairs), spicate, ebracteate; peduncle 0.5 –1.5 cm long, puberulous; spike 6 –12 cm long;
flowers numerous; tepals c. 1.5 mm long, sparsely puberulous; ovary c. 1 mm long,
style c. 0.5 mm long, stigmas 1.5 – 2 mm long. Infructescences green to pale yellow.
Endocarp body ellipsoid to ovoid, c. 0.7 mm long. — Fig. 1; Map 1.
Distribution — India (Assam), Sikkim, Bhutan, South China (incl. Hainan), Myan-
mar, Cambodia, Thailand; in Malesia: Sumatra, Java.
Habitat — Montane and submontane forest, often on volcanic soil, at altitudes be-
tween 400 and 2500 m.
Uses — Its timber is useful for house construction.
Note — The features, including the long pistillate inflorescences, show similarities
to M. insignis Bureau, a species of humid montane forests in the Neotropics.
BLEEKRODEA
Bleekrodea Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 87; Benth. & Hook.f., Gen. Pl. 3 (1880)
358, ‘Bleekrodia’; Leandri, Fl. Madagasc. fam. 55 (1952) 5; C.C. Berg, Proc. Kon. Ned. Akad.
Wetensch. C, 91 (1988) 359. — Streblus Lour. sect. Bleekrodea (Blume) Corner, Gard. Bull. Sin-
gapore 19 (1962) 216; Phytomorphology 25 (1975) 6, t. 5, 6.
Shrubs or treelets, monoecious, unarmed. Leaves distichous; lamina pinnately veined;

stipules free, lateral. Inflorescences bisexual (or unisexual), short-branched cymose to
subcapitate, sometimes subinvolucrate. Staminate flowers several to one (or lacking),

pedicellate; tepals 4 or 5, valvate in the bud; stamens 4 (or 5), inflexed in the bud. Pis
tillate flowers 1 or 2 in the centre of the inflorescence (or lacking), (sub)sessile; tepals
4, connate, forming a tubular, dentate perianth; ovary free, stigmas 2, equal. Fruiting
perianth enlarged, slightly fleshy, pink; fruit free, a dehiscent drupe, whitish, endocarp
thinly crustaceous; seed without endosperm, cotyledons thick, flat, unequal to subequal,
radicle short.
Distribution — The genus comprises two species, one in Malesia and the other one
in Madagascar.
Morphology — These two species are rather similar in their features.
The inflorescences are bisexual with 1 or 2 pistillate flowers or unisexual. The stami-
nate inflorescences contain several to many flowers and are in structure similar to the
pistillate ones. The pistillate inflorescences are uniflorous and bear a few bracts.
The structure of the inflorescences of Streblus asper and S. tonkinensis (see p. 42)
resemble that of Bleekrodea, but this genus is distinct by the tubular perianth of the
pistillate flower and the presence of uncinate hairs.
1. Bleekrodea insignis Blume

Bleekrodea insignis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 88, f. 28. — Streblus insignis
(Blume) Corner, Gard. Bull. Singapore 19 (1962) 219; Phytomorphology 25 (1975) t. 5.
Streblus malayanus Corner, Phytomorphology 25 (1975) 11, t. 6. — Bleekrodea malayana (Corner)
C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 359.
Treelet. Leafy twigs 1– 2 mm thick, minutely puberulous, partly with uncinate hairs.
Leaves distichous; lamina elliptic to subobovate, 5 –14(– 21) by 2 – 5(–7.5) cm, sub
coriaceous, apex acuminate to subcaudate, base cuneate, margin crenate-dentate; upper
and lower surface sparsely puberulous on the main veins, smooth; lateral veins 6 –10,
tertiary venation reticulate (to subscalariform); petiole 0.3 –1 cm long, minutely puberu-
lous, partly with uncinate hairs; stipules 0.3 – 0.6 cm long, puberulous, subpersistent.
Inflorescences axillary, solitary, bisexual or staminate and 0.7– 2 cm wide or pistillate
and uniflorous; peduncle 0.5 –1.2 cm long, minutely puberulous; staminate flowers:
pedicel 0.5 –1.5 mm long, perianth 1.5 mm long, puberulous, stamens 2 – 2.5 mm long,
anthers c. 1 mm long; pistillate flower(s): perianth 3 – 4 mm, in fruit up to c. 6 mm
long, densely puberulous, style 3 – 6 mm long, stigmas 3 – 8 mm long; bracts ovate to
narrowly ovate, 2 – 3 mm long, puberulous, bracteoles narrowly ovate to subulate, 3 – 5
mm long, puberulous. Endocarp body c. 1 cm long. — Fig. 2.
Distribution — Malay Peninsula (Kelantan: Gua Musang), Borneo.
Habitat — Lowland forest, in the Malay Peninsula on limestone.
Notes — 1. The collection from Peninsular Malaysia described as Streblus malaya
nus and an additional collection from the same locality largely match material known
from Borneo.
2. The variation in the composition of the inflorescences resembles that of the Mada-
gascan species, Bleekrodea madagascariensis Blume (see Berg, Bull. Jard. Bot. Belg.
47 (1977) 367).
1.5 cm
5 mm
c
f
e
4 cm
1.5 cm
a
b 8 mm
d
Fig. 2. Bleekrodea insignis Blume. a. Leaf; b. ripe fruit in section: 1 = utricular perianth, 2 = exocarp,
3 = membranous testa, 4 = cotyledon, 5 = radicle; c. embryo in section: 1 = radicle; d. inflorescence
at anthesis of the staminate flowers, one pistillate flower in fruit; e, f. staminate flowers (a–f: Jaheri
1541).
BROUSSONETIA
Broussonetia L’Hér. ex Vent., Tabl. Règn. Vég. 3 (1799) 547, nom. cons., non Gomez Ortega, Nov.
Rar. Pl. Hort. Matr. Descr. (1798) 61 (Legum.); Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856)
85; Bureau in A.DC., Prodr. 17 (1873) 223; Baill., Hist. Pl. 6 (1875) 192; Benth. & Hook.f., Gen.
Pl. 3 (1880) 361; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 76; Boerl., Handl. Fl. Ned.
Ind. 3 (1900) 316; Corner, Gard. Bull. Singapore 19 (1962) 233; H.L. Li, Woody Fl. Taiwan (1963)
112; Ohwi, Fl. Japan, new ed. (1965) 383; C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 355.
Papyrius Lam., Tabl. Encycl. (1798) t. 762.
Allaeanthus Thwaites, Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 302; Bureau in A.DC., Prodr. 17
(1873) 222; Benth. & Hook.f., Gen. Pl. 3 (1880) 361; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3,
1 (1888) 74 (‘Allacanthus’); Boerl., Handl. Fl. Ned. Ind. 3 (1900) 317. — Broussonetia L’Hér. ex
Vent. sect. Allaeanthus (Thwaites) Corner, Gard. Bull. Singapore 19 (1962) 234.
Smithiodendron Hu, Sunyatsenia 3 (1936) 106.
Trees (or climbers), dioecious, unarmed, deciduous, shoot apices shed, with (small)
axillary scaled resting buds. Leaves distichous, spirally arranged or (sub)opposite;
lamina pinnately veined; stipules free, semi-amplexicaul. Inflorescences unisexual, usu-

ally solitary in the leaf axils or on leafless nodes at the base of fertile twigs, bracteate.
Staminate inflorescences spicate, sometimes subcapitate, with a rather distinct abaxial
sterile strip; perianth 4-parted, segments valvate in the bud; stamens 4, inflexed in
the bud. Pistillate inflorescences globose-capitate, sometimes elongated; perianth un-
equally 2 – 4-lobed to 2 – 4-dentate; ovary free, stigmas 2, one of them strongly reduced;
interfloral bracts longer than the perianth, their upper parts cohering by entangling
hairs. Fruiting perianth enlarged, membranous; fruit usually short-stipitate and slightly
compressed, exocarp fleshy, for the greater part rather thinly so, thicker at the base and
on the seed-bearing side; endocarp crustaceous, with a thickened crest-like part at the
apex and a small woody plug towards the hilum; seed with endosperm, embryo curved,
cotyledons unequal or almost equal, subconduplicate to almost flat, straight or the apex
reflexed, not enclosing the long radicle, or if the cotyledons unequal, then the larger
cotyledon more or less clasping the smaller one; bracts more or less fused at the base
and hardened at the apex in fruit.
Distribution — A genus of eight species, seven in tropical to subtropical Asia, and
one in Madagascar.
Subdivision — Two sections (Broussonetia and Allaeanthus) have been recognised
by Corner (Gard. Bull. Singapore 19 (1962) 234). As he indicated, the differences be-
tween these sections are very small, and are not maintained in the present treatment.
KEY TO THE SPECIES
1a. Leaves spirally arranged or subopposite; lamina scabrous above. — Widespread,

in cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. B. papyrifera
b. Leaves distichous; lamina smooth above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Tree; margin of the lamina entire. — Philippines, Celebes . . . . . . . 2. B. luzonica
b. Climber; margin of the lamina dentate. — Sumatra . . . . . . . . . . . . . . 1. B. kurzii
1. Broussonetia kurzii (Hook.f.) Corner

Broussonetia kurzii (Hook.f.) Corner, Gard. Bull. Singapore 19 (1962) 234. — Allaeanthus kurzii
Hook.f., Fl. Brit. India 5 (1888) 490; Gagnep., Fl. Indo-Chine 5 (1928) 721; Kanjilal & A.C. Das,
Fl. Assam 4 (1940) 275.
Malaisia tortuosa (non Blume) Kurz, Forest Fl. Burma 2 (1877) 466.
Climber, deciduous. Leafy twigs 1– 3 mm thick, appressedly puberulous. Leaves

distichous; lamina narrowly elliptic to elliptic or subovate to ovate, 4 –12 by 2.5 – 6 cm,
± asymmetric, chartaceous, apex acuminate to subacute, base rounded to subcordate,
margin crenate- to serrate-dentate; upper surface sparsely appressedly puberulous on the
main veins, smooth; lower surface sparsely appressedly puberulous on the veins; midrib
impressed above, lateral veins 5 – 8 pairs, tertiary venation scalariform; petiole 0.6 –1.5
cm long, sparsely appressedly puberulous; stipules 0.3 – 0.9 cm long, ovate, sparsely
minutely puberulous, caducous or subpersistent. Staminate inflorescences clustered on
short-shoots or on the base of leafy twigs, spicate; peduncle 0.3 – 2 cm long, minutely
puberulous; spike (0.5 –)1– 5 cm long; perianth 1.5 – 2 mm long, minutely puberulous;
stamens 2.5 – 3 mm long, anthers 0.6 – 0.8 mm long; bracts ovate to linear, puberulous.
Pistillate inflorescences solitary on up to 4 cm long leafless branchlets, or at the base
of leafy twigs, capitate; peduncle 0.3 –1 cm long, minutely puberulous to tomentellous;
head globose, 0.2 – 0.4 cm diam.; perianth c. 0.5 mm long, 4-dentate; ovary c. 0.3 mm
long, stigma 7–10 mm long; interfloral bracts clavate to subpeltate, c. 0.7 mm long,
densely puberulous to tomentellous, bracts subtending the head ovate, 0.5 –1 mm long,
sparsely minutely puberulous. Infructescences globose, 0.5 – 0.8 cm diameter.
Distribution — India (Assam), Myanmar, Thailand; in Malesia: N Sumatra (Atjeh,
known by a single collection).
Habitat — Lowland rain forest (edges).
Note — The infructescences are eaten by gibbons.
2. Broussonetia luzonica (Blanco) Bureau

Broussonetia luzonica (Blanco) Bureau in A.DC., Prodr. 17 (1873) 224; Corner, Gard. Bull. Singapore
19 (1962) 235. — Morus luzonica Blanco, Fl. Filip. (1837) 703. — Allaeanthus luzonicus (Blanco)
Fern.-Vill., Nov. App. (1880) 198; S. Vidal, Sin. Gen. Pl. Leños Filip. (1883) 39, t. 86 f. A; Phan.
Cuming. Philipp. (1885) 145; Revis. Pl. Vasc. Filip. (1886) 250; Merr., Sp. Blancoan. (1918) 122;
Enum. Philipp. Flow. Pl. 2 (1923) 37.
Broussonetia luzoniensis Blanco, Fl. Filip. ed. 2 (1845) 488; ed. 3, 3 (1879) 107, t. 278.
Allaeanthus glaber Warb. in Perkins, Fragm. Fl. Philipp. 3 (1905) 166. — Allaeanthus luzonicus
(Blanco) Fern.-Vill. var. glaber (Warb.) Merr., Enum. Philipp. Flow. Pl. 2 (1923) 37. — Broussone
tia luzonica (Blanco) Bureau var. glabra (Warb.) Corner, Gard. Bull. Singapore 19 (1962) 235.
Fig. 3. Broussonetia luzonica (Blanco) Bureau.

a. Twig with staminate inflorescences; b, c. en- 2 cm
docarp bodies; d. fruit in section: 1 = radicle,
2 = membranous testa, 3 = endocarp, 4 = exo-
carp; e. embryo (a: Herb. Lugd. Bat. 908.186-
118; b – e: Bur. Sci. 42294).
5 mm
3 mm
e d
5 mm
b a c 5 mm
Tree up to 10 m tall (or more?), deciduous. Leafy twigs 1.5 – 2.5 mm thick, sparsely
to densely puberulous to (sub)tomentellous. Leaves distichous; lamina subovate to
ovate, 6 –17 by 2.5 – 6 cm, ± asymmetric, chartaceous, apex acuminate to subacute, base
rounded to obtuse or to subcordate, margin entire; upper surface sparsely puberulous,
smooth; lower surface sparsely to densely puberulous, to tomentose on the veins; midrib
impressed above, lateral veins 7–12 pairs, tertiary venation scalariform; petiole 0.6 –1.5
cm long, sparsely to densely puberulous to tomentose; stipules narrowly ovate, 0.3 –
0.9 cm long, puberulous, ribbed, caducous. Staminate inflorescences axillary, solitary,
spicate; peduncle 1–1.5 cm long, puberulous to tomentellous; spike 5 – 26 cm long;
perianth 1–1.5 mm long, densely minutely puberulous; stamens c. 2 mm long, anthers
c. 0.7 mm long; bracts ovate to narrowly elliptic, 0.5 –1 mm long, minutely puberulous.
Pistillate inflorescences axillary, solitary, capitate; peduncle 0.6 –1 cm long, puberulous
to tomentellous; head globose, 0.4 – 0.7 cm diam.; perianth c. 1 mm long, 4-dentate;
ovary c. 0.5 mm long, stigma 5 –15 mm long; interfloral bracts clavate to subpeltate,
1–1.3 mm long, densely puberulous to tomentellous. Infructescences globose, 1.5 – 2
cm diameter.— Fig. 3.
Distribution — Philippines, Celebes.
Habitat — Lowland forest
3. Broussonetia papyrifera (L.) Vent.

Broussonetia papyrifera (L.) Vent., Tabl. Règn. Vég. 3 (1799) 547; Blume, Mus. Bot. 2 (1856) 85;
Miq., Ann. Mus. Bot. Lugduno-Batavi 2 (1866) 198; Bureau in A.DC., Prodr. 17 (1873) 224; Hook.
f., Fl. Brit. India 5 (1888) 490; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 3; Koord.,
Exkurs.-Fl. Java 2 (1912) 87; Atlas Baumart. Java (1924) t. 746; Gagnep., Fl. Indo-Chine 5 (1928)
717, f. 88; F. Brown, Bull. Bish. Mus. 130 (1935) 33; T.S. Liu, Ill. Lign. Pl. Taiwan 2 (1962) 708;
H.L. Li, Woody Fl. Taiwan (1963) 113; Ohwi, Fl. Japan, new ed. (1965) 384; Backer & Bakh.f.,
Fl. Java 2 (1965) 16; Barker, Curtis’s Bot. Mag. 19 (2002) 8, cum t.; C.C. Berg, Prosea 17 (2003)
91, cum t. — Morus papyrifera L., Sp. Pl. (1753) 986.
Broussonetia papyrifera (L.) Vent. forma sumatrana Miq., Fl. Ned. Ind., Eerst Bijv. (1861) 416. —
Broussonetia papyrifera (L.) Vent. var. sumatrana (Miq.) Bureau in DC., Prodr. 17 (1873) 224.
Smithiodendron artocarpoideum Hu, Sunyatsenia 3 (1936) 106.
Tree up to 35 m tall, deciduous. Leafy twigs 1.5 – 3 mm thick, subtomentose to hirtel-

lous to puberulous. Leaves spirally arranged or (sub)opposite; lamina ovate to cordiform
or to elliptic, 5 – 20 by 4 –12 cm, entire or lobate, ± asymmetric, chartaceous, apex acu-
minate to subacute, base cordate to rounded to subcuneate, margin crenate- to serrate-
dentate; upper surface hispidulous, scabrous; lower surface ± densely puberulous, to
subtomentose on the veins; midrib impressed above, lateral veins 5 – 9 pairs, tertiary
venation scalariform; petiole (1–)2 – 9(–15) cm long, subtomentose; stipules 0.5 –1.5
cm long, ovate, puberulous, ± ribbed, caducous. Staminate inflorescences axillary or
below the leaves, solitary or clustered on short-shoots, spicate; peduncle 1– 2.5 cm long,
puberulous to tomentellous; spike 3 –10 cm long; perianth 1.5 – 2 mm long, puberulous;
stamens 3 – 3.5 mm long, anthers c. 0.8 mm long; bracts subulate, 1.5 – 2.5 mm long,
puberulous. Pistillate inflorescences axillary or below the leaves, solitary, capitate;
peduncle 0.3 –1.5 cm long, puberulous to tomentellous; head globose, 1–1.2 cm diam.;
perianth c. 1 mm long, 4-dentate; ovary 0.5 mm long, stigma 7–10 mm long; interflo-
4 mm
5 mm
5 mm
2.5 mm
h i g
b
c
10 mm
a d e
2 cm 1 cm
Fig. 4. Broussonetia papyrifera (L.) Vent. a. Pistillate inflorescence in section at anthesis; b. pistillate
flower at anthesis, with bracts, the perianth cut open; c. base of pistillate flower; d. fruit-containing
perianths with elongate bracts; e. fruit with the enlarged utricular perianth cut open; f. stalked drupe in
section; g. embryo; h, i. endocarp bodies (material from Cambridge Botanic Garden).
ral bracts clavate to subpeltate, 1–1.5 mm long, densely puberulous to tomentellous.

Infructescences (sub)globose, 2 – 2.5 cm diam.; endocarp body ovoid, 2 – 2.5 mm long.
— Fig. 4.
Distribution — India (Assam), China (incl. Taiwan), Indochina, Japan (introduced in
the Ryukyu Islands), Myanmar, Thailand, Polynesia; in Malesia: introduced in Sumatra,
Java, Philippines, Celebes, Lesser Sunda Islands (Flores, Timor, Alor, Wetar), Moluc-
cas, New Guinea.
Habitat — Lowland and highlands, in particular in parts with seasonal climate.
Uses — The bark is used to make cloth and paper (see Barker 2002 and Berg 2003).
FATOUA
Fatoua Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509; Bureau in A.DC., Prodr. 17 (1873) 255;
Baill., Hist. Pl. 6 (1875) 197; Benth. & Hook.f., Gen. Pl. 3 (1880) 358; Engl. in Engl. & Prantl,
Nat. Pflanzenfam. 3, 1 (1888) 71; C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 370.
Subshrubs or herbs, monoecious, latex absent. Leaves distichous; lamina pinnately

veined, with cystolith hairs beneath; stipules free, lateral. Inflorescences bisexual (or
unisexual), in pairs or solitary in the axils of the leaves, short-branched cymose or spi-
cate, abaxially without flowers. Staminate flowers sessile or short-pedicellate; perianth
4-parted, segments valvate; stamens 4, inflexed in the bud, Pistillate flowers sessile;
perianth 4-parted, segments valvate; ovary free, style lateral, one of the stigmas strongly
reduced. Fruiting perianth enlarged, not fleshy; exocarp thick, white fleshy and de-
hiscent (probably) ejecting the endocarp body; seed with endosperm, embryo curved,
cotyledons equal and flat, radicle long.
Distribution — The genus comprises two species, one extending from Japan to New
Caledonia and the other only known from Madagascar.
Morphology — The genus shows similarities to Bleekrodea in the bisexual inflo-
rescences and the dehiscent drupes. Moreover, in F. villosa the inflorescence is short-
branched cymose like in Bleekrodea. However, in the Madagascan species, F. mada
gascariensis Leandri (1948), the inflorescence is racemose to spicate.
An anatomical study by Leandri, Mem. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 1
B (1948) 4 revealed the absence of lactiferous elements in the Madagascan species.
1. Fatoua villosa (Murray) Nakai

Fatoua villosa (Murray) Nakai, Bot. Mag. (Tokyo) 41 (1927) 516. — Urtica villosa Murray, Syst. Veg.
ed. 14 (July 1784) 851; Thunb., Fl. Jap. (Aug. 1784) 70.
Urtica japonica Thunb., Fl. Jap. (Oct. 1784) 70, non L.f. 1781.
Fatoua pilosa Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509; Gaudich., Voy. Bonite, Bot. (1844)
t. 84 f. 1; Bureau in A.DC., Prodr. 17 (1873) 256; Benth., Fl. Australia 6 (1873) 182; Engl. in Engl.
& Prantl, Nat. Pflanzenfam. 3, 1 (1888) 71, f. 50A, B; Warb., Bot. Jahrb. Syst. 13 (1891) 294; F.B.
Forbes & Hemsl., J. Linn. Soc., Bot. 26 (1894) 454; Koord., Exkurs.-Fl. Java 2 (1912) 83; Merr.,
Enum. Philipp. Flow. Pl. 2 (1923) 36; Koord., Atlas Baumart. Java (1924) t. 743; Gagnep., Fl. Indo-
Chine 5 (1928) 715; Backer & Bakh.f., Fl. Java 2 (1965) 14.
Fatoua aspera Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 510; Miq., Fl. Ned. Ind. 1, 2 (1859) 282;
Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 192.
Fatoua cordata Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 510.
Fatoua lanceolata Decne., Nouv. Ann. Mus. Hist. Nat. 3 (1834) 492; Gaudich., Voy. Bonite, Bot. (1844)
t. 84 f. 3; Miq., Fl. Ned. Ind. 1, 2 (1859) 282.
Urtica manillensis Walp., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19, Suppl. 1 (1843) 423.
Fatoua subcordata Gaudich., Voy. Bonite, Bot. (1844) t. 84 f. 2.
Fleurya glechomaefolia Miq. in Zoll., Syst. Verz. 2 (1854) 106. — Fatoua pilosa Gaudich. (var. cor
data) subvar. glechomaefolia (Miq.) Bureau in A.DC., Prodr. 17 (1873) 257.
Fleurya globulifera Miq. in Zoll., Syst. Verz. 2 (1854) 106. — Fatoua pilosa Gaudich. (var. cordata)
subvar. globulifera (Miq.) Bureau in A.DC., Prodr. 17 (1873) 257.
Fatoua japonica Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) t. 38.
Fatoua pilosa Gaudich. var. lanceolata Bureau in A.DC., Prodr. 17 (1873) 257.
Herb or subshrub up to 1 m tall, usually branched, erect; roots yellow. Leafy twigs
0.5 – 2 mm thick, minutely puberulous and with longer mostly curved to uncinate
hairs. Leaves distichous; lamina cordiform to ovate (to narrowly ovate), 1– 6(– 9) by
0.5 – 4(– 5) cm, membranaceous, apex acuminate to (sub)acute, base cordate to trun-
cate to rounded (to subcuneate), margin dentate to serrate or to crenate; upper surface
hispidulous to puberulous to hirtellous, scabrous to smooth; lower surface minutely
puberulous, on the veins the short indumentum intermixed with longer hairs; lateral
veins 3 –7(–10), running into the margin or faintly loop-connected, tertiary venation
subscalariform to reticulate; petiole 0.5 – 4 cm long, minutely puberulous and with
longer curved to uncinate hairs; stipules 0.3 – 0.5 cm long, puberulous to hirtellous,
subpersistent. Inflorescences subcapitate, 0.3 –1 cm diam.; peduncle 0.5 – 2.5 cm long,

minutely puberulous; staminate flowers: tepals c. 1 mm long, puberulous, stamens
c. 1.5 mm long, anthers c. 0.5 mm long; pistillate flowers: tepals c. 1 mm long, style
2 – 3 mm long. Endocarp body c. 0.8 by 0.6 mm.
Distribution — China (incl. Taiwan), Japan, Thailand, Indochina, Australia, Solo-
mon Islands, New Caledonia; in Malesia: Java (also Madura and Bawean), Philippines,
Celebes (also Buton), Lesser Sunda Islands, Moluccas (Halmahera, Banda, Ceram,
Tanimbar Islands, Kai Islands), New Guinea (also Bismarck Archipelago).
Habitat — Forest edges, seacoasts, light secondary forest, waste places, on calcare-
ous soil; at altitudes up to c. 1200 m.
Notes — 1. There are no clear indications that the species is in Malesia represented
by perennials (Fatoua pilosa?) and annuals (F. villosa?) see Zhou Zhekun & H.G.
Gilbert, Fl. China 5 (2003) 22.
2. The species is very variable as with regard to habit, indumentum, and leaf char-
acters, but varieties cannot be distinguished.
3. The venation of the lamina is craspedodromous rather than brochidodromous as
characteristic for the family.
MACLURA
Maclura Nutt., Gen. N. Amer. Pl. 2 (1818) 233, nom. cons.; Blume, Ann. Mus. Bot. Lugduno-Batavi
2 (1856) 81; Bureau in A.DC., Prodr. 17 (1873) 226; Baill., Hist. Pl. 6 (1875) 193; Benth. & Hook.
f., Gen. Pl. 3 (1880) 363; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 74; Lemée, Dict.
Gen. Pl. Phan. 4 (1932) 230; Corner, Gard. Bull. Singapore 19 (1962) 235; C.C. Berg, Proc. Kon.
Ned. Akad. Wetensch. C, 91 (1986) 243; Fl. Neotrop. Monogr. 83 (2001) 53.
Vanieria Lour., Fl. Cochinch. 1 (1790) 564.
Ioxylon Raf., Amer. Monthly Mag. & Crit. Rev. 2 (1817) 118; 4 (1818) 188 (‘Toxylon’),
Chlorophora Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509. — Maclura Nutt. sect. Chlorophora
(Gaudich.) Corner, Gard. Bull. Singapore 19 (1962) 236.
Fusticus Raf., New Fl. Am. 3 (1836) 43.
Sukaminea Raf., New Fl. Am. 3 (1836) 44.
Cudrania Rumph. ex Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 122, nom. cons.; Bureau, Ann. Sci.
Nat., Bot. sér. 5, 11 (1869) 377; Bureau in A.DC., Prodr. 17 (1873) 285; Baill., Hist. Pl. 6 (1875)
202; Benth. & Hook.f., Gen. Pl. 3 (1880) 374; Hook.f., Fl. Brit. India 5 (1888) 538; Engl. in Engl.
& Prantl, Nat. Pflanzenfam. 3, 1 (1888) 82; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 329 (‘Cudravia’);
Renner, Bot. Jahrb. Syst. 39 (1907) 361. — Cudranus Rumph. [Herb. Amboin. 5 (1747) 22] ex
Miq., Fl. Ned. Ind. 1, 2 (1859) 290. — Maclura Nutt. sect. Cudrania (Trécul) Corner, Gard. Bull.
Singapore 19 (1962) 237.
Plecospermum Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 124; Corner, Gard. Bull. Singapore 19
(1962) 241. — Maclura Nutt. sect. Plecospermum (Trécul) C.C. Berg, Proc. Kon. Ned. Akad.
Wetensch. C, 91 (1986) 245.
Cardiogyne Bureau in A.DC., Prodr. 17 (1873) 232; C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 359.
— Maclura Nutt. sect. Cardiogyne (Bureau) Corner, Gard. Bull. Singapore 19 (1962) 237.
x Maclucudrania André, Rev. Hort. 77 (1905) 363.
Climbers, trees or shrubs, dioecious, armed with straight to curved, reduced branch-
lets ending in a spinose tip (sometimes only in juvenile specimens). Leaves distichous
or spirally arranged; lamina pinnately veined; stipules free or fused, lateral to semi-
amplexicaul, often very small. Inflorescences usually solitary, in the leaf axils or on
short-shoots, often yellow (dye-containing) glands embedded in the bracts and/or

the (fruiting) perianths. Staminate inflorescences globose-(sub)capitate, spicate, or
(sub)racemose; perianth with 4, free or basally connate, tepals, decussate, imbricate
in the bud; stamens straight or inflexed in the bud. Pistillate inflorescences globose-
capitate; flowers free or connate; tepals free or connate, decussate, imbricate in the bud;
ovary free or the lower part adnate to the perianth, stigmas 2, mostly strongly unequal
in length, or 1. Fruiting perianth enlarged, more or less fleshy; fruit free or adnate to
the perianth, when free slightly drupaceous, endocarp crustaceous; seed small to rather
large, endosperm present (but scarce) or absent, embryo various, mostly with (rather)
thin, folded or flat, cotyledons and a long radicle.
Distribution — The genus comprises eleven species of which ten occur in the tropics
(seven in Asia, one in Africa, and two in the Neotropics) and one in North America.
Subdivision — The genus can be subdivided into five sections (Berg 1986): Cardio
gyne (one species in Africa and another in the Neotropics), Chlorophora (one species
in the Neotropics), Cudrania (four species in Asia), Maclura (a single species in North
America), and Plecospermum (two species in Asia). The species of sect. Cudrania do
not have the urticaceous type of stamen and are exceptional in the tribe Moreae.
The species occurring in the Malesian region represent sect. Cudrania ser. Connatae
Corner (Gard. Bull. Singapore 19 (1962) 238). It differs from the three other species of
this section in the basally connate pistillate flowers with one stigma or two unequally
long ones.
Dispersal — The orange-coloured pulpy infructescences are dispersed by birds
(Ridley, Dispersal of plants throughout the world, 1930).
Section Cudrania
Maclura Nutt. sect. Cudrania (Trécul) Corner, Gard. Bull. Singapore 19 (1962) 237.
Climbers or shrubs. Staminate inflorescences globose-capitate; stamens straight in

the bud. Pistillate flowers free or connate; tepals and bracts accumulating yellow dye.
1. Maclura cochinchinensis (Lour.) Corner

Maclura cochinchinensis (Lour.) Corner, Gard. Bull. Singapore 19 (1962) 239; Backer & Bakh.f.,
Fl. Java 2 (1965) 17; Steenis, Mt Fl. Java (1972) pl. 30 –7. — Vanieria cochinchinensis Lour., Fl.
Cochinch. (1790) 564; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 39; Nakai, Bot. Mag. (Tokyo) 41
(1927) 515. — Procris cochinchinensis (Lour.) Spreng., Syst. Veg. 3 (1826) 846. — Cudrania
cochinchinensis (Lour.) Kudô & Masam., Ann. Rep. Taihoku Bot. Gard. 2 (1932) 27; Merr., Comm.
Fl. Cochinch. (1935) 134; Kudô & Masam., Trans. Nat. Hist. Soc. Taiwan (1938) 292; Backer,
Blumea 6 (1948) 302; T.S. Liu, Ill. Lign. Pl. Taiwan 2 (1962) 709; H.L. Li, Woody Fl. Taiwan
(1963) 113, f. 36; Ohwi, Fl. Japan, new ed. (1965) 384.
Vanieria alternifolia Stokes, Bot. Mat. Med. 4 (1812) 381.
Trophis spinosa Blume, Bijdr. (1825) 489, non Willd. 1806, nec Roxb. 1832. — Cudrania spinosa
(Blume) Hochr., Bull. New York Bot. Gard. 6 (1910) 271; Koord., Exkurs.-Fl. Java 2 (1912) 90.
Morus tinctoria (non L. ) Blanco, Fl. Filip. (1837) 704. — Broussonetia tinctoria Blanco, Fl. Filip. ed.
2 (1845) 488; ed. 3, 3 (1879) 108, t. 418, non (L.) Kunth 1817. — Maclura gerontogea Siebold &
Zucc., Abh. Akad. München 4, 3 (1846) 220; Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 84.
— Vanieria cochinchinensis Lour. var. gerontogea (Siebold & Zucc.) Nakai, Bot. Mag. (Tokyo)
41 (1927) 516. — Cudrania cochinchinensis (Lour.) Kudô & Masam. var. gerontogea (Siebold &
Zucc.) Kudô & Masam., Ann. Rep. Taihoku Bot. Gard. 2 (1932) 27.
Cudrania pubescens Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 125; Miq., Fl. Ned. Ind. 1, 2 (1859)
290, ‘Cudranus pubescens’; Kurz, Forest Fl. Burma 2 (1877) 435; Koord., Exkurs.-Fl. Java 2
(1912) 91; Gagnep., Fl. Indo-Chine 5 (1928) 727. — Vanieria pubescens (Trécul) Chun, J. Arnold
Arbor. 8 (1927) 21. — Maclura cochinchinensis (Lour.) Corner var. pubescens (Trécul) Corner,
Gard. Bull. Singapore 19 (1962) 239. — Maclura pubescens (Trécul) Z.K. Zhou & M.G. Gilbert,
Fl. China 5 (2003) 36.
Plecospermum spinosum Trécul var. javanensis Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 125.
Cudrania javanensis Trécul var. indica Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 123.
Cudrania obovata Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 126; Miq., Fl. Ned. Ind. 1, 2 (1859) 290,
‘Cudranus obovatus’; Gagnep., Fl. Indo-Chine 5 (1928) 730.
Cudrania sumatrana Miq., Pl. Jungh. (1851) 44; Fl. Ned. Ind. 1, 2 (1859) 291, ‘Cudranus sumatranus’.
Cudrania acuminata Miq. in Zoll., Syst. Verz. 2 (1854) 90 nomen, 96 descr.; Miq., Fl. Ned. Ind. 1, 2
(1859) 291, ‘Cudranus acuminatus’.
Plecospermum cuneifolium Thwaites, Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 303.
Maclura timorensis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 54.
Maclura javanica Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 83; Miq., Fl. Ned. Ind. 1, 2 (1859)
280. — Cudrania javanensis Rumph. ex Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 123, t. 3,
f. 76 – 85; Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1960; Miq., Fl. Ned. Ind. 1, 2 (1859) 290; Bureau,
Ann. Sci. Nat., Bot. sér. 5, 11 (1869) 378; Benth., Fl. Australia 6 (1873) 179; Hook.f., Fl. Brit. India
5 (1888) 538; in Trimen, Fl. Ceylon 4 (1898) 98; K. Schum. & Lauterb., Fl. Schutzgeb. Südsee
(1901) 266; Merr., Interpr. Herb. Amboin. (1917) 189; Sp. Blancoan. (1918) 123, ‘javensis’; Koord.,
Exkurs.-Fl. Java Atlas (1924) t. 750; Ridl., Fl. Malay Penins. 3 (1924) 356; Gagnep., Fl. Indo-
Chine 5 (1928) 726; Diels, Bot. Jahrb. Syst. 67 (1935) 174. — Cudranus rumphii Thwaites, Enum.
Pl. Zeyl. (1861) 262. — Cudranus spinosus Kuntze, Revis. Gen. Pl. 2 (1891) 625. — [Cudranus
bimanus sive javanus Rumph. Herb. Amboin. 5 (1747) 22, t. 15, f. 2.]
Maclura amboinensis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 84; Corner, Gard. Bull. Singa
pore 19 (1962) 239; Backer & Bakh. f., Fl. Java 2 (1965)17. — Cudranus amboinensis (Blume)
Miq., Fl. Ned. Ind. 1, 2 (1859) 290; Kurz, Fl. Burma 2 (1877) 434, ‘Cudrania’; Warb., Bot. Jahrb.
Syst. 13 (1891) 294. — [Cudranus amboinicus Rumph., Herb. Amboin. 5 (1747) 22, t. 15 f. 1, t.
16.]
Cudrania rectispina Hance, J. Bot. 14 (1876) 365; F.B. Forbes & Hemsl., J. Linn. Soc., Bot. 26 (1899)
470.
Cudrania grandifolia Merr., Philipp. J. Sci. 18 (1921) 52. — Vanieria grandifolia (Merr.) Merr., Enum.
Philipp. Flow. Pl. 2 (1923) 39.
Cudrania cambodiana Gagnep., Bull. Soc. Bot. France 72 (1925) 808; Fl. Indo-Chine 5 (1928) 729,
t. 89: 9 –14, 90: 1– 3.
Maclura amboinensis Blume var. paucinervia Corner, Gard. Bull. Singapore 19 (1962) 239.
Shrub, often much-branched with long scrambling sprays, or climber, with up to

4 cm long straight or curved thorns. Leafy twigs 1.5 – 4 mm thick, brownish to whitish
puberulous to subglabrous, often drying brown with numerous whitish lenticels. Leaves
spirally arranged to distichous; lamina elliptic to (sub)obovate to narrowly elliptic,
(0.5 –)4 – 8(–12) by (0.3 –)2 – 5 cm, subcoriaceous to chartaceous, apex (short-)acu-
minate to subacute (to apiculate or to rounded), base obtuse to rounded or to cuneate,
margin entire; upper surface glabrous; lower surface (very) sparsely minutely puberu-
lous on the (main) veins to glabrous; midrib impressed above, lateral veins 6 – 9(–14)
pairs, tertiary venation reticulate or subscalariform; petiole 0.3 –1.5(– 2) cm long,
(sub)glabrous; stipules 0.2 – 0.3 cm long, puberulous, caducous or subpersistent. Stami
nate inflorescences axillary, in pairs or solitary, capitate; peduncle 0.3 –1 cm long,
5 mm
5 mm
f g
3 cm
2 cm l
e i
2 mm
1 cm
4 cm
6 cm
d
1 cm
j
5 mm
a b c k h
Fig. 5. Maclura cochinchinensis (Lour.) Corner. a. Leaf showing the transverse intercostal veins; b. leaf
showing the absence of transverse intercostals; c. pistillate inflorescence at anthesis in section; d. pistil-
late flower at anthesis, showing basally connate flowers; e. infructescence in section; f. embryo with
the cotyledon-folds cut away to show the long radicle; g. embryo in transverse section: 1, 2 = radicle;
h. sapling leaf; i. leaf of a very young plant; j. staminate inflorescence at anthesis, in section, the anthers
not exserted; k. open staminate flower in section; 1. mature stamen (a: Herb. Lugd. Bat. 908.157-618;
b: Reinwardt s.n.; c – f: Bünnemeijer 8110; h, i: Bakhuizen van den Brink 2949; j – l; Walsh 300).
puberulous to tomentellous; head globose, 0.4 –1 cm diam.; tepals c. 1.5 mm long,

minutely puberulous; stamens c. 1 mm long, anthers c. 0.5 mm long; bracts basally
attached, 0.5 –1 mm long, minutely puberulous. Pistillate inflorescences axillary, in
pairs or solitary, capitate; peduncle 0.4 –1.2 cm long, puberulous to tomentellous; head
globose, 0.4 – 0.6 cm diam.; flowers basally connate; perianth c. 1 mm long, densely
white puberulous to tomentellous; ovary c. 1 mm long, style 0.5 mm long, stigmas 2 – 3
mm long, twisted; bracts basally attached, 0.5 –1 cm long, in fruit up to 3 mm long,
sparsely to densely puberulous. Infructescences globose, 1.5 – 2 cm diam., yellow to
orange to red. — Fig. 5.
Distribution — Sri Lanka, India, Bhutan, Sikkim, Japan, Australia, New Caledonia;
in Malesia: throughout.
Habitat — Lowland forest, chiefly forest edges, by mangrove forest, riverbanks, and
in secondary forest, rarely in high forest, at altitudes up to 1800 m.
Uses — The infructescences are edible and the wood is used for dyeing silk yellow
or green.
STREBLUS
Streblus Lour., Fl. Cochinch. (1790) 615; Bureau in A.DC., Prodr. 17 (1873) 218; Baill., Hist. Pl. 6
(1875) 195; Benth. & Hook.f., Gen. Pl. 3 (1880) 359; Engl. in Engl. & Prantl, Nat. Pflanzenfam.
3, 1 (1888) 78; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 314; Merr., Comm. Fl. Cochinch. (1935) 134;
Corner, Gard. Bull. Singapore 19 (1962) 215; Blumea 18 (1970) 393; Phytomorphology 25 (1975) 1;
C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 356; Kochummen, Tree Fl. Malaya 3
(1978) 167; Go, Tree Fl. Sabah & Sarawak 3 (2000) 329; C.C. Berg, Blumea 50 (2005) 547.
Achymus Juss., Dict. Sci. Nat. 1, Suppl. (1816) 31; Poir., Dict. Sci. Nat. 51 (1827) 91, ‘Achimus’.
Epicarpurus Blume, Bijdr. (1825) 488.
Albrandia Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509; D. Dietr., Syn. Pl. 5 (1852) 280, ‘Albra
dia’.
Calius Blanco, Fl. Filip. (1837) 698.
Ampalis Bojer, Hort. Maurit. (1837) 291; Corner, Gard. Bull. Singapore 19 (1962) 214; C.C. Berg,
Bull. Jard. Bot. Belg. 47 (1977) 343. — Streblus Lour. sect. Ampalis (Bojer) C.C. Berg, Proc. Kon.
Ned. Akad. Wetensch. C, 91 (1988) 356.
Taxotrophis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 77; Hutch., Bull. Misc. Inform. Kew (1918)
147. — Streblus Lour. sect. Taxotrophis (Blume) Corner, Gard. Bull. Singapore 19 (1962) 218.
Paratrophis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 81; Cheeseman, Man. New Zealand Fl.
(1906) 631. — Streblus Lour. sect. Paratrophis (Blume) Corner, Gard. Bull. Singapore 19 (1962)
216.
Sloetia Teijsm. & Binn. ex Kurz, J. Linn. Soc., Bot. 8 (1864) 168. — Streblus Lour. sect. Sloetia (Teijsm.
& Binn. ex Kurz) Corner, Gard. Bull. Singapore 19 (1962) 218.
Pseudomorus Bureau, Ann. Sci. Nat., Bot. sér. 5, 11 (1869) 372; Skottsb., Acta Horti Gothob. 15 (1944)
347; Stearn, J. Arnold Arbor. 28 (1947) 426.
Diplocos Bureau in A.DC., Prodr. 17 (1873) 215.
Phyllochlamys Bureau in A.DC., Prodr. 17 (1873) 217. — Streblus Lour. sect. Phyllochlamys (Bureau)
Corner, Gard. Bull. Singapore 19 (1962) 217.
Pseudostreblus Bureau in A.DC., Prodr. 17 (1873) 219; Hook.f., Fl. Brit. India 5 (1888) 487. — Stre
blus Lour. sect. Pseudostreblus (Bureau) Corner, Gard. Bull. Singapore 19 (1962) 217; Phytomor-
phology 25 (1975) 6, t. 3C.
Pachtytrophe Bureau in A.DC., Prodr. 17 (1873) 234; Corner, Gard. Bull. Singapore 19 (1962) 214;
C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 339.
Uromorus Bureau in A.DC., Prodr. 17 (1873) 236.
Pseudotrophis Warb., Bot. Jahrb. Syst. 13 (1890) 291. — Streblus Lour. sect. Pseudotrophis (Warb.)
Sloetiopsis Engl., Bot. Jahrb. Syst. 39 (1907) 573.
Teonongia Stapf in Hook., Icon. Pl. 30 (1913) t. 2947; Gagnep., Fl. Indo-Chine 5 (1928) 710.
Neosloetiopsis Engl., Bot. Jahrb. Syst. 51 (1914) 426.
Dimerocarpus Gagnep., Bull. Mus. Hist. Nat. (Paris) 27 (1921) 441.
Chevalierodendron Leroy, Compt. Rend. Hebd. Séances Acad. Sci. 227 (1948) 146.
Trees or shrubs, dioecious or monoecious, unarmed or armed with thorns, uncinate

hairs often present. Leaves distichous or sometimes spirally arranged; lamina pin-
nately veined, with cystoliths above and beneath, only beneath, or absent; stipules
free or sometimes fused, lateral or sometimes almost fully amplexicaul. Inflorescences
unisexual or sometimes bisexual, racemose, spicate or subcapitate, bracteate; stami-
nate inflorescences multi- to pluriflorous; pistillate inflorescences multi- to uniflorous.
Staminate flowers with 3, 4, or 5 tepals, imbricate or valvate in the bud; stamens 3,
4, or 5, inflexed in the bud. Pistillate flowers 4-merous; tepals (almost) free, more or
less unequal in size; ovary free, stigmas 2, equal. Fruiting perianth enlarged or not,
somewhat fleshy or not, mostly greenish; fruit free, drupaceous, dehiscent and whitish
or indehiscent and yellow, orange, red, or blackish, endocarp (thinly) crustaceous; seed
without endosperm, cotyledons folded or not, equal or unequal, radicle short or long.
Distribution — The genus comprises 23 species in the tropics of the Old World.
Subdivision — By reduction of many former genera to sections, Corner (1962) has
widened the circumscription of the genus considerably. Corner (1962, 1970) recognised
eight sections in the genus. A more simplified subdivision has been proposed by Berg
(1988), who excluded sect. Bleekrodea, because of the structure of the inflorescences
and the utriculate perianth of the pistillate flower, and included three African–Mada-
gascan species. The sections recognised are: section Ampalis (comprising two Madagas-
can species), section Paratrophis (comprising eight Australasian–Asian species), the
monotypic sections Protostreblus and Sloetia, and the Asian–African section Streblus
(comprising eight Asian and one African species). Section Streblus, in this narrow sense,
differs from the other sections in the enlarged fruiting perianths, and includes four
non-aculeate species with mostly subcoriaceous laminas and dentate margins. Section
Taxotrophis comprises six aculeate species and section Pseudostreblus (Bureau) Corner
the non-aculeate species with coriaceous laminas and entire margins (the latter contains
only S. indicus (Bureau) Corner (1962), ranging from Myanmar to northern Thailand,
South China (incl. Hainan) and Cambodia (see Berg 2005)). — Map 2.
Morphology — In several features including inflorescences, leaves, ecological dif-
ferentiation, and number of species Streblus shows clear similarities to the neotropical
genus Sorocea A. St.-Hil. In several species of sect. Streblus thorns are formed. In some
species the drupes are dehiscent.
Ecology — Several species of sect. Streblus are associated with relatively dry and/or
seasonal conditions, whereas some species of sect. Paratrophis are associated with
southern warm-temperate conditions, or (S. glaber) partly with montane conditions.
Map 2. Distribution of Streblus Lour. sect. Paratrophis (Blume) Corner in dotted line (the westward
extension of sect. Paratrophis is the submontane distribution of S. glaber in central and western Ma-
laysia), and the other sections in broken line. Distribution of S. perakensis in closed line.
Relationships — The genus shows parallel morphological differentiation to the ge-

nus Trophis and the similarities suggest that the two genera are closely related. The
free tepals of ± unequal size of the pistillate flower, not becoming fleshy in Streblus,
versus the fused ones of equal size, usually becoming fleshy in Trophis, is the major
differentiating character. In all Trophis species, except for the peculiar Central American
T. involucrata W.C. Burger, the perianth becomes enlarged to fully enclosing the fruit,
and turns red, purplish, or orange at maturity.
Akad. Wetensch. C, 91 (1988) 345 – 362. — Berg, C.C., Fl. Neotrop. Monogr. 83 (2001) 32. — Cor-
ner, E.J.H., The classification of Moraceae. Gard. Bull. Singapore 19 (1962) 187– 252. — Corner,
E.J.H., New species of Streblus and Ficus (Moraceae). Blumea 18 (1970) 393 – 411.
KEY TO THE SECTIONS
1a. Leaves spirally arranged; petiole 1.5 – 6 cm long; upper surface of the lamina ± sca-
brous and base of the lamina cordate to rounded . . . . . . . . . . Sect. Protostreblus
b. Leaves distichous, petiole up to 1.5 cm long, or if longer (up to 3.5 cm), then upper
surface of the lamina smooth and base of the lamina cuneate to rounded . . . . . . . 2
2a. Inflorescences, bisexual or staminate, spicate; staminate flowers 3-merous . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sect. Sloetia
b. Inflorescences unisexual, spicate, racemose, (sub)capitate, or (pistillate ones) uni-
florous, if bisexual, then the inflorescence (sub)capitate with a single pistillate
flower; staminate flowers 4-merous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Pistillate inflorescences spicate to subcapitate, usually with 2 or more flowers
and the perianth not enlarged in fruit; staminate inflorescences spicate and the
peduncle usually at least 0.4 cm long; bracts peltate or basally attached . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sect. Paratrophis
b. Pistillate inflorescences racemose with 2 –15 flowers, or uniflorous, the perianth in
fruit enlarged; staminate inflorescences spicate and the peduncle short (0 – 0.4 cm
long) or capitate; bracts basally attached . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Plants without thorns, often monoecious; lamina mostly subcoriaceous and the mar-
gin dentate; staminate inflorescences capitate; pistillate inflorescences uniflorous;
tepals of staminate flowers imbricate; tepals of pistillate flowers in fruit reflexed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sect. Streblus
b. Plants (always or sometimes) with thorns, dioecious; lamina mostly coriaceous and
the margin and/or acumen spinulose; pistillate inflorescences (sub)racemose or
uniflorous; staminate inflorescences spicate (to subcapitate); tepals of staminate
flowers valvate; tepals of pistillate flowers not reflexed in fruit . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sect. Taxotrophis
Section Streblus
Streblus Lour. sect. Streblus; Corner, Gard. Bull. Singapore 19 (1962) 218; C.C. Berg, Proc. Kon. Ned.
Akad. Wetensch. C, 91 (1988) 357.
Achymus Juss., Dict. Sci. Nat. 1, Suppl. (1816) 31.
Epicarpurus Blume, Bijdr. (1825) 488.
Albrandia Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509.

Calius Blanco, Fl. Filip. (1837) 698.
Sloetiopsis Engl., Bot. Jahrb. Syst. 39 (1907) 573; C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 363.
Teonongia Stapf, Hooker’s Icon. Pl. 30 (1911) t. 2947.
Neosloetiopsis Engl., Bot. Jahrb. Syst. 51 (1914); C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 363.
Diplothorax Gagnep., Bull. Soc. Bot. France 75 (1928) 98.
Trees or shrubs, monoecious or dioecious, without spines. Leaves distichous; lamina

elliptic, subcoriaceous to chartaceous or to coriaceous, margin dentate; petiole short;
stipules free, lateral. Inflorescences bisexual or unisexual, capitate or cymose, pistillate
ones usually uniflorous, bracts basally attached; staminate flowers 4-merous, tepals
imbricate; fruiting perianth more or less enlarged, reflexed (or not). Drupe dehiscent
or not.
Distribution — This section comprises four species, two of them in the Asian main-
land and western Malesia, one in Celebes and the fourth in Africa.
Morphology — Monoecy occurs in Streblus asper and, more pronouncedly so in
S. tonkinensis (Eberh. & Dubard) Corner (incl. S. monoicus Gagnep. = S. asper var.
monoicus (Gagnep.) Corner), a species from Laos and Vietnam. The African species,
S. usambarensis (Engl.) C.C. Berg (Bull. Jard. Bot. Belg. 47 (1977) 364; Proc. Kon.
Ned. Akad. Wetensch. C, 91 (1988) 357) is distinct in the spicate staminate inflores-
cences and the fruiting perianth remaining patent; the latter feature may prove to occur
in S. celebensis as well.
KEY TO THE SPECIES
1a. Indumentum whitish; lamina ± scabrous above; lateral veins 4 –7(– 8) pairs. —
Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. S. asper
b. Indumentum predominantly dark brown; lamina glabrous above; lateral veins (7–)
8 –10 pairs. — Celebes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. celebensis
1. Streblus asper Lour.

Streblus asper Lour., Fl. Cochinch. 1 (1790) 615; Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 79,
f. 33; Hook.f., Fl. Brit. India 5 (1888) 489; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 7;
Koord., Exkurs.-Fl. Java 2 (1912) 88; Merr., Sp. Blancoan. (1918) 123; Enum. Philipp. Flow. Pl.
2 (1923) 38; Koord., Exkurs.-Fl. Java, Atlas (1924) f. 747; Gagnep., Fl. Indo-Chine 5 (1928) 712,
f. 86: 10 –16, 87: 1– 2; Diels, Bot. Jahrb. Syst. 67 (1935) 172; Merr., Comm. Fl. Cochinch. (1935)
134; Alston, Kandy Fl. (1938) 35, F. 184; Corner, Wayside Trees Malaya ed.1 (1940) 692, t. 255;
Worth., Ceylon Trees (1959) pl. 424; Corner, Gard. Bull. Singapore 19 (1962) 228; Backer & Bakh.
f., Fl. Java 2 (1965) 16; Corner, Phytomorphology 25 (1975) 3, t. 1B, 3D; Fl. Ceylon 1.2 (1977)
156, t. 28; Wayside Trees Malaya ed. 3 (1988) 555, t. 168.
Trophis aspera Retz., Observ. Bot. 5 (1788) 30; Roxb., Fl. Ind., ed. Carey 3 (1832) 761; Wight, J. Bot.
1 (1834) 62, t. 121.
Trophis cochinchinensis Poir., Encycl. 8 (1808) 123.
Trophis aculeata Roth, Nov. Pl. Sp. (1821) 868.
Epicarpurus orientalis Blume, Bijdr. (1825) 488; Hassk., Pl. Jav. Rar. (1848) 197; Miq., Pl. Jungh.
(1851) 42; Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1961; Zoll., Syst. Verz. (1854) 89.
Calius lactescens Blanco, Fl. Filip. (1837) 698; ed. 3, 3 (1879) 1103, t. 171. — Streblus lactescens
(Blanco) Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 80.
Achymus pallens Sol. ex Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 79.
Cudrania crenata C.H. Wright, J. Linn. Soc., Bot. 26 (1899) 469; Merr., Lingnan Sci. J. 16 (1937) 187.
— Vanieria crenata (C.H. Wright) Chun, J. Arnold Arbor. 8 (1927) 21.
Diplothorax tonkinensis Gagnep., Bull. Soc. Bot. France 75 (1928) 98.
Shrub or tree up to 10(–15) m tall, much-branched, branches often drooping; lower

ones (often?) prostrate, monoecious or dioecious. Leafy twigs 1– 2 mm thick, whitish
puberulous. Leaves distichous; lamina elliptic to obovate or subobovate, 1– 8(–13) by
0.5 – 3.5(– 6.5) cm, coriaceous to subcoriaceous, apex (sub)acuminate to acute, base
rounded to subcordate to obtuse, margin crenate to dentate; upper surface sparsely
whitish hispidulous to puberulous, ± scabrous (or almost smooth); lower surface whit-
ish hispidulous to puberulous, ± scabrous; midrib ± prominent in the lower part of the
lamina, ± impressed in the upper part, lateral veins 4 –7(– 8) pairs, tertiary venation
reticulate (to subscalariform); petiole 0.1– 0.3(– 0.5) cm long, whitish puberulous;
stipules 0.3 – 0.5 cm long, puberulous, caducous (or subpersistent). Staminate inflo
rescences axillary, in pairs or solitary, capitate; peduncle 0.2 –1.5 cm long, sparsely
minutely whitish puberulous; head 0.4 –1 cm diam.; flowers 4 –15; perianth 1.5 – 2 mm
long, whitish minutely puberulous; stamens 2 – 2.5 mm long, anthers c. 1 mm long;
bracts few, basally attached, 0.5 – 2 mm long, sparsely minutely puberulous. Pistillate
inflorescences axillary, in pairs or solitary, uniflorous (or biflorous); peduncle 0.4 –1.5
cm long, minutely whitish puberulous; tepals 2 – 2.5 mm long, minutely puberulous,
in fruit 5 – 8 mm long, reflexed; ovary c. 1 mm long, style c. 1 mm long, elongating up
to 3 mm, stigmas 2 – 4 mm long, elongating up to 12 mm; bracts few, basally attached,
0.5 – 2 mm long, sparsely minutely puberulous. Drupe (sub)globose, 0.6 – 0.8 cm long,
indehiscent, yellow to orange at maturity. — Fig. 6; Map 3.
Distribution — Sri Lanka, India, Bhutan, Bangladesh, Myanmar, Andaman Islands,
Nicobar Islands, Thailand, South China (incl. Hainan), Indochina; in Malesia: Sumatra
Map 3. Distribution of Streblus asper Lour.

e
1 cm f 5 mm
2 cm
i
b
h
1 cm
d 1 cm
2 cm
1 cm
a c g
Fig. 6. Streblus asper Lour. a. Pistillate inflorescence, one-flowered, at anthesis; b. the same in section;
c. with nearly ripe fruit; d. ripe fruit; e. ripe fruit in section: 1 = side of the large cotyledon, 2 = radi-
cle, 3 = plumule, 4 = small cotyledon, 5 = exocarp, 6 = endocarp, 7 = membranous testa, 8 = inner
face of the large cotyledon; f. embryo in end-view, showing the radicle (1) and large cotyledon;
g. staminate inflorescence in section; h. staminate flower; i. staminate flower-bud in section (material
from Penang).
(Atjeh, Medan), Malay Peninsula (Penang, Kedah, Perlis, Kelantan), Java (also Ma-
dura and Kangean Island), Philippines, Celebes, Lesser Sunda Islands (Bali, Lombok,
Sumbawa, Flores, Sumba), Moluccas (Ambon).
Habitat — Strongly seasonal monsoon climate, chiefly in secondary forest, by rice-
fields and in open places; at low altitudes.
Uses — The fruits are edible, sweet and the leaves can be used as cattle fodder.
2. Streblus celebensis C.C. Berg

Streblus celebensis C.C. Berg, Blumea 50 (2005) 547.
Shrub, 1 m tall, without spines, dioecious. Leafy twigs 1– 2 mm thick, puberulous,

mainly with dark brown appressed hairs. Leaves distichous; lamina subcoriaceous,
(sub)obovate, 6 –15 by 2 –7 cm, apex acuminate to subcaudate, base obtuse, margin

coarsely crenate-dentate towards the apex; upper surface (subglabrous) smooth; lower
surface sparsely appressed puberulous with dark brown appressed hairs, scabridulous;
midrib prominent or prominent in a groove above, lateral veins (7–)8 –10 pairs, tertiary
venation subscalariform to reticulate, slightly prominent; petiole 0.3 – 0.7 cm long, c. 1
mm thick, puberulous with dark brown appressed hairs; stipules 0.1– 0.2 cm long, sub-
persistent (or caducous). Staminate inflorescences axillary, solitary, subsessile, subcapi-
tate with 3 – 5 flowers; tepals 4, valvate in the bud, c. 1.5 mm long, outside appressed
puberulous with dark brown hairs, the margin white tomentellous; stamens 4, inflexed
in the bud, anthers c. 1 mm long, pistillode obconical, c. 0.5 mm long; bracts basally
attached ovate 1– 2 mm long, appressed puberulous. Pistillate inflorescences axillary,
solitary, uniflorous; peduncle c. 0.2 cm long; tepals 4, decussate imbricate, c. 1.5 mm
long, outside appressed puberulous with dark brown hairs, margin white ciliolate; ovary
c. 1 mm long, style c. 3 mm long, stigmas 2, filiform, 6 – 8 mm long; bracts few, basally
attached, ovate, c. 0.2 mm long, appressedly puberulous; tepals enlarging and probably
enveloping the fruit at full maturity.
Distribution — Celebes (only known from Batudaka Island).
Habitat — Humid forest at low altitudes.
Section Paratrophis
Streblus Lour. sect. Paratrophis (Blume) Corner, Gard. Bull. Singapore 19 (1962) 216; Phytomor
phology 25 (1975) 1; C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 357. — Paratrophis
Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 81.
Uromorus Bureau in A.DC., Prodr. 17 (1873) 236.
Chevalierodendron Leroy, Compt. Rend. Hebd. Séances Acad. Sci. 227 (1948) 146.
Trees, unarmed, dioecious. Leaves distichous; lamina elliptic to narrowly elliptic;

petiole short; stipules free, lateral. Inflorescences unisexual, spicate, bracts peltate or
basally attached; staminate flowers 4-merous, tepals valvate in the bud; pistillate flow-
ers usually 2 or more; fruiting perianth not or hardly enlarged. Drupe indehiscent, red
or blackish at maturity.
Distribution & Delimitation — This section comprises eight species and it is dis-
tinguishable from sect. Protostreblus by the distichous arrangement of the leaves, and
from the other Asian – Australasian sections by the tepals of the pistillate flower, which
are not or only slightly enlarged in fruit. This section is distinctly concentrated in the
eastern part of the range of the genus. One of the species, Streblus pendulinus, extends
just into the Malesian region. Only S. glaber is widespread in this region and occurs also
in Australia (Queensland). The other five are elements of the Pacific region: Streblus
anthropophagarum (Seem.) Corner, incl. S. solomonenis Corner (from the Solomon
Islands to Marquesas Islands), S. banksii (Cheeseman) C.J. Webb (New Zealand J.
Bot. 25 (1987) 136) in New Zealand, S. heterophyllus (Blume) Corner (1962) in New
Zealand, S. sclerophyllus Corner (Blumea 18 (1970) 399, t. 4) in New Caledonia, S.
smithii (Cheeseman) Corner (1962) in New Zealand, and S. tahitensis (Nadeaud) Corner
(1962) in Tahiti.
KEY TO THE SPECIES
1a. Midrib of the lamina ± prominent above, margin mostly subentire, lamina (sub)
coriaceous. — Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. S. glaber
b. Midrib of the lamina impressed above, margin serrate to subspinulose dentate,
lamina chartaceous to subcoriaceous. — New Guinea . . . . . . . . 4. S. pendulinus
3. Streblus glaber (Merr.) Corner

Streblus glaber (Merr.) Corner, Gard. Bull. Singapore 19 (1962) 221; Phytomorphology 25 (1975)
2, t. 3B; Go, Tree Fl. Sabah & Sarawak 3 (2000) 331, t. 16. — Gironniera glabra Merr., Philipp.
J. Sci., 1, Suppl. (1906) 42; Enum. Philipp. Flow. Pl. 2 (1923) 35. — Paratrophis glabra (Merr.)
Steenis, J. Bot. 72 (1934) 8. — Chevalierodendron glabrum (Merr.) Leroy, Compt. Rend. Hebd.
Séances Acad. Sci. 227 (1948) 146.
Tree up to 40 m tall or shrub, dioecious. Leafy twigs 0.5 – 2 mm thick, (sub)glabrous

or whitish minutely puberulous. Leaves distichous; lamina narrowly elliptic to elliptic
to subovate to narrowly ovate (or to subrhombic or linear), 1.5 –16 by 0.5 – 4.5 cm,
(sub)coriaceous, apex acuminate to subcaudate, base cuneate to almost rounded (or
to subattenuate), margin subentire (or dentate); upper surface glabrous; lower surface
puberulous on the main veins or glabrous, often with minute red or red-brown pluricel-
lular trichomes; midrib ± prominent above, lateral veins 5 –18 pairs (or if the lamina is
linear, then up to 30 pairs), tertiary venation reticulate; petiole 0.3 –1.7 cm long, sparsely
whitish appressedly puberulous; stipules 0.2 – 0.5 cm long, minutely whitish appress-
edly puberulous, caducous. Staminate inflorescences axillary, up to 5 together, spicate
(or uniflorous); peduncle 0.3 –1.5 cm long, sparsely minutely puberulous; spike 0.8–2
cm or up to 3 cm long; flowers (1–)3 –15 or up to 30; perianth 1– 2 mm long, minutely
puberulous or only ciliolate, stamens 2.5 – 3 mm long, anthers 1–1.2 mm long; bracts
basally attached, 0.3 –1 mm long, sparsely minutely puberulous. Pistillate inflores
cences axillary, up to 5 together, short-spicate to subcapitate or uniflorous; peduncle
0.5 – 2 cm long, minutely puberulous; flowers 1– 3(– 9); tepals 1–1.5 mm long, ciliolate
or subglabrous, in fruit hardly enlarged; ovary 1.5 – 2 mm long, stigmas c. 2 mm long;
bracts basally attached, 0.3 –1 mm long, ciliolate or subglabrous. Fruit ellipsoid to
ovoid, 0.8 –1.2 cm, dark red to purple at maturity.
Notes — 1. Two subspecies can be recognised in the Malesian region: the typical
subspecies widespread in lowland to submontane and a montane one confined to east-
ern New Guinea. The main differentiating morphological character is the presence or
absence of teeth in the margin of the lamina. However, in Sabah subsp. glaber, which
has normally an entire lamina margin, can have a dentate one. There is an overlap of
the altitudinal ranges of the two subspecies and transitional forms are encountered.
2. The number of inflorescences in the leaf axils is more than the common pair, as
short-shoots are formed in the leaf axils with solitary or paired inflorescences on the
nodes. This axillary leafless shoot sometimes becomes longer and may bear up to 12
inflorescences.
3. The Australian material of S. glaber (Merr.) Corner, Gard. Bull. Singapore 19
(1962) 221 (— Paratrophis australiana C.T. White, Contr. Arnold Arbor. 4 (1933) 15)
occurring at altitudes up to 1000 m, differs from both subspecies in the usually crenate-
dentate margin of the lamina. It was referred to S. glaber var. australianus (C.T. White)
Corner (Gard. Bull. Singapore 19 (1962) 221) and is currently recognised as subsp.
australianus (C.T. White) C.C. Berg (Blumea 50 (2005) 548).
KEY TO THE MALESIAN SUBSPECIES
1a. Lamina dentate, at least 2 teeth in the upper part of the lamina. — E New Guinea
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. urophyllus
b. Lamina entire (sometimes dentate in Borneo). — Widespread . . . a. subsp. glaber
1 cm
1 cm
1 cm
f
2 cm
b 3 mm
1 cm
1 cm
a c e g
Fig. 7. Streblus glaber (Merr.) Corner subsp. glaber. a, b. Leaves showing the absence of transverse
intercostal veins; c. many-flowered pistillate inflorescence; d. ovary in section; e. few-flowered pistil-
late inflorescence; f. fruits; g. staminate inflorescence; h. staminate flower (a: Clemens 32132; b: NGF
6924; c, d: Elmer 22014; e: Eyma 4863; f: SFN 26946; g, h: Clemens 29321).
a. subsp. glaber
Aphananthe negrosensis Elmer, Leafl. Philipp. Bot. 2 (1909) 575.
Pseudostreblus caudatus Ridl., J. Fed. Malay States Mus. 6 (1915) 54; Fl. Malay Penins. 3 (1924) 324.
Streblus laevifolius Diels, Bot. Jahrb. Syst. 67 (1935) 171.
Tree up to 25(– 40) m tall. Lamina elliptic or to narrowly ovate, (1.5 –)3 –16 by
(0.5 –)1– 4.5 cm, margin of the lamina usually entire, but sometimes dentate in the upper
part of the lamina (in Borneo: Sabah); lateral veins 7–18 pairs. — Fig. 7.
Distribution — Solomon Islands; in Malesia: Sumatra, Malay Peninsula, Borneo
(mainly Sabah), Philippines (Luzon, Mindoro, Negros, Cebu), Celebes, Moluccas (Aru
Islands), New Guinea.
Habitat — Montane forest, sometimes on limestone or in relatively dry areas; mostly
at altitudes between 700 and 2500 m, rarely down to sea level.
Note — In the Philippines, the inflorescences are usually longer (up to 3 cm long)
and both staminate and pistillate inflorescences tend to have more flowers, often more
than 15 or 3, respectively.
1 cm
2 cm
4 cm
Fig. 8. Streblus glaber (Merr.) Corner subsp. uro

phyllus (Diels) C.C. Berg. a – d. Leaf-variation;
e. fruit in section; f, g. embryos to show the obli
quity (a: J.C. Sanders 879; b: Robbins 199; c, d:
a b c Pullen 310; e–g: Hoogland & Pullen 5529).
b. subsp. urophyllus (Diels) C.C. Berg

Streblus glaber (Merr.) Corner subsp. urophyllus (Diels) C.C. Berg, Blumea 50 (2005) 548; Corner,
Gard. Bull. Singapore 19 (1962) 225. — Streblus urophyllus Diels, Bot. Jahrb. Syst. 67 (1935)
172.
Streblus urophyllus Diels var. salicifolius Corner, Gard. Bull. Singapore. 19 (1962) 225.
Shrub or tree up to 15 m tall. Lamina narrowly elliptic to elliptic to subovate to nar-

rowly ovate (or to subrhombic) or linear, 2 – 8(–15) by 0.5 – 3(– 4.5) cm, margin of the
lamina dentate to denticulate, at least two teeth in the upper part of the lamina; lateral
veins 5 –10 pairs, but if the lamina linear (willow-leaf shaped) then up to 30 pairs.
— Fig. 8.
Distribution — New Guinea (eastern).
Habitat — Primary (or secondary) forest at altitudes between (1600 –)2400 and 3000 m,
occasionally on limestone.
Note — For this subspecies a form with linear laminas and (therefore) numerous
lateral veins has been described as var. salicifolius.
4. Streblus pendulinus (Endl.) F. Muell.

Streblus pendulinus (Endl.) F. Muell., Fragm. Phyt. Australiae 6 (1868) 192; Corner, Gard. Bull.
Singapore 19 (1962) 222; Blumea 18 (1970) 397, t. 2. — Morus pendulina Endl., Prod. Fl. Norfolk.
(1833) 40. — Pseudomorus pendulina (Endl.) Stearn, J. Arnold Arbor. 28 (1947) 427.— Pseudo
morus brunoniana (Endl.) Bureau var. pendulina (Endl.) Bureau, Ann. Sci. Nat., Bot. sér. 4, 11
(1869) 373.
Morus brunoniana Endl., Atakta Bot. 4 (1835) t. 32. — Streblus brunonianus (Endl.) F. Muell., Fragm.
Phyt. Australiae 6 (1868) 192. — Pseudomorus brunoniana (Endl.) Bureau, Ann. Sci. Nat., Bot.
sér. 4, 11 (1869) 373; Rock, Indig. Trees Haw. Isl. (1913) 114. — Pseudomorus brunoniana (Endl.)
Bureau var. australiana Bureau, Ann. Sci. Nat., Bot. sér. 4, 11 (1869) 373.
Pseudomorus brunoniana (Endl.) Bureau var. obtusata Bureau, Ann. Sci. Nat., Bot. sér. 4, 11 (1869)
373.
Pseudomorus sandwicensis O. Deg., Fl. Hawaiiensis (1938) fam. 96. — Pseudomorus brunoniana
(Endl.) Bureau var. sandwicensis (O. Deg.) Skottsb., Acta Horti Gothob. 15 (1944) 347.
Shrub or tree up to 13 m tall, dioecious (or monoecious). Leafy twigs 1– 2 mm thick,

(minutely) whitish puberulous. Leaves distichous; lamina elliptic to (sub)ovate, (1.5 –)
4 –10(–14) by (0.5 –)1.5 – 4.5(– 6.5) cm, chartaceous to subcoriaceous, apex acuminate
to (sub)acute to subobtuse, base subcordate to cuneate, margin serrate to subspinulose-
dentate, sometimes lobate; upper surface glabrous and smooth (or scabrous); lower
surface whitish puberulous on the main veins, scabridulous; midrib impressed above,
lateral veins (4 –)7–10(–14) pairs, tertiary venation reticulate; petiole 0.2 –1.1 cm long,
(minutely) whitish puberulous; stipules 0.2 – 0.4 cm long, whitish puberulous, caducous.
Staminate inflorescences axillary, solitary or in pairs, spicate; peduncle (0.2 –)0.4 – 0.7
(– 2.5) cm long, (minutely) puberulous; spike 2 –18 cm long; flowers numerous; perianth
1–1.5 mm long, sparsely minutely puberulous, stamens c. 2.5 mm long, anthers c. 0.8
mm long; bracts peltate, c. 0.5 mm diam., ciliolate. Pistillate inflorescences axillary,
solitary or in pairs, short-spicate to subcapitate; peduncle 0.2 – 0.8 cm long; flowers
2 – 9; tepals 1–1.5 mm long, ciliolate, in fruit hardly enlarged; ovary 1.5 – 2 mm long,
stigmas 2 – 3 mm long; bracts (sub)peltate, 0.5 –1 mm diam., ciliolate. Fruit ellipsoid
to ovoid to subglobose, 0.6 –1 cm long, yellow to reddish to brownish or to purple at

maturity.
Distribution — Australia (Queensland, New South Wales), and in the Pacific: New
Hebrides, New Caledonia, Norfolk Island, Fiji, Rapa, Hawaii, Guam, Saipan, Rota; in
Malesia: New Guinea (eastern).
Habitat — Lowland forest (as on limestone hills), on seashore, and in swamps, often
under seasonal conditions.
Note — Collection W. Moi 149 from Papua New Guinea, Morobe Province, deviates
from the other collections from New Guinea in being monoecious and having laminas
scabrous above.
Section Protostreblus
Streblus Lour. sect. Protostreblus Corner, Blumea (1970) 393; C.C. Berg, Proc. Kon. Ned. Akad.
Wetensch. C, 91 (1988) 358.
Trees, unarmed, dioecious. Leaves spirally arranged; lamina ovate to subcordiform;

petiole long; stipules free, lateral. Inflorescences unisexual, bracts basally attached.
Staminate inflorescences unknown. Pistillate inflorescences spicate, multiflorous; fruit-
ing perianth hardly enlarged. Drupe indehiscent, yellow to orange (or red?) at matu-
rity.
Delimitation — This section comprises a single species and is distinguished by the
spirally arranged leaves with long petioles.
5. Streblus ascendens Corner

Streblus ascendens Corner, Blumea 18 (1970) 395, t. 1; Phytomorphology 25 (1975) 2.
Tree up to 23 m tall, with buttresses up to 1.5 m high, dioecious. Leafy twigs 2 – 4

mm thick, whitish to brownish puberulous. Leaves spirally arranged; lamina ovate
to subcordiform, 9 –14(– 24) by 5 – 8(–15) cm, subcoriaceous, apex acuminate, base
(sub)cordate to rounded, margin subentire to irregularly crenate-dentate; upper sur-
face sparsely whitish puberulous, ± scabrous; lower surface whitish puberulous to sub
tomentose; midrib slightly impressed above, lateral veins 3 or 4 pairs, tertiary venation
scalariform; petiole 1.5 – 6 cm long, whitish appressedly puberulous; stipules 0.4 – 0.6
cm long, ± densely whitish to yellowish appressedly puberulous, caducous. Staminate
inflorescences unknown. Pistillate inflorescences axillary, solitary, spicate; peduncle
0.7–1.2 cm long, minutely puberulous; flowers numerous; tepals c. 2 mm long, minutely
puberulous, in fruit up to 4 mm long; ovary c. 1.5 mm long, stigmas 1.5 – 2 mm long;
bracts basally attached, 0.3 –1 mm long, minutely puberulous. Drupe obovoid, c. 0.7
cm long when dry, yellow to orange (to red?) at maturity. — Fig. 9.
Distribution — Solomon Islands (Kolombangara); in Malesia: New Guinea (Mano
kwari).
Habitat — Lowland forest.
5 mm
b
2 cm
a
1 mm
2.5 mm
d
Fig. 9. Streblus ascendens Corner. a. Leafy twig with pistillate inflorescence; b. pistillate spike in sec-
tion; c. pistillate flowers; d. ripe fruit (endocarp and hilar plug hatched) (all: BSIP 8624).
Section SLOETIA
Streblus Lour. sect. Sloetia (Teijsm. & Binn. ex Kurz) Corner, Gard. Bull. Singapore 19 (1962) 218;
Phytomorphology 25 (1975) 4; C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 358. —
Sloetia Teijsm. & Binn. ex Kurz, J. Linn. Soc., Bot. 8 (1864) 168.
Trees, unarmed, monoecious. Leaves distichous; lamina elliptic; petiole short; stip-
ules free, nearly fully amplexicaul. Inflorescences bisexual or staminate, spicate, bracts
peltate; staminate flowers numerous, 3-merous, tepals imbricate in the bud; pistillate
flowers 0 – 4; fruiting perianth enlarged. Drupe dehiscent.
Delimitation — This section comprises a single species and is distinguished by the
spicate, mostly bisexual inflorescences and 3-merous staminate flowers.
6. Streblus elongatus (Miq.) Corner

Streblus elongatus (Miq.) Corner, Gard. Bull. Singapore 19 (1962) 227; Phytomorphology 25 (1975)
4, t. 1D, 3E; Kochummen, Tree Fl. Malaya 3 (1978) 168, t. 11; Go, Tree Fl. Sabah & Sarawak
3 (2000) 330. — Artocarpus elongatus Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 419. — Sloetia
elongata (Miq.) Koord., Exkurs.-Fl. Java 2 (1912) 90, Koord., Atlas Baumart. Java (1924) t. 749;
Burkill, Dict. Econ. Prod. Malay Penins. (1935) 2035; Corner, Wayside Trees Malaya ed.1 (1940)
691, t. 254.
Sloetia sideroxylon Teijsm. & Binn. ex Kurz, J. Linn. Soc., Bot. 8 (1864) 168, t. 13; Bureau in A.DC.,
Prodr. 17 (1873) 257; Hook.f., Fl. Brit. India 5 (1888) 493; Foxw., Malayan Forest Rec. 3 (1927)
129.
Sloetia sideroxylon Teijsm. & Binn. ex Kurz var. brevipes Bureau in A.DC., Prodr. 17 (1873) 258.
Sloetia penangiana Oliv., Hooker’s Icon. Pl. (1886) t. 1531; Hook.f., Fl. Brit. India 5 (1888) 493.
Sloetia wallichii King ex Hook.f., Fl. Brit. India 5 (1888) 493.
Tree up to 35 m tall, often flowering as shrub, monoecious. Leafy twigs 1.5 – 2.5 mm
thick, sparsely whitish puberulous with uncinate hairs. Leaves distichous; lamina nar-
rowly elliptic to subobovate to subovate to narrowly ovate (or to elliptic), 6 – 25(– 38)
by 1.5 – 8(–16) cm, usually ± asymmetric, coriaceous to subcoriaceous, apex acuminate,
base cuneate to obtuse, margin entire (or towards the apex sparsely denticulate); upper
surface glabrous; lower surface sparsely whitish puberulous on the main veins, smooth
(or scabridulous); midrib prominent above, lateral veins 10 – 20 pairs, tertiary venation
(sub)scalariform; petiole 0.2 –1.8 cm long, sparsely whitish puberulous to subglabrous;
stipules 0.5 –1(–1.5) cm long, sparsely minutely whitish puberulous to subglabrous,
caducous (or subpersistent). Inflorescences axillary, solitary or in pairs, spicate, stami-
nate or bisexual and protogynous; peduncle 0.2 –1 cm long, sparsely puberulous to sub
glabrous; spike 3 – 20 cm long; staminate flowers numerous; perianth 3- (or 4-)merous,
1–1.2 mm long, minutely puberulous; stamens 3 (or 4), 2 – 2.5 mm long, anthers 0.6 – 0.8
mm long; pistillate flowers 0 – 4; tepals 4, c. 1.5 mm long, ciliolate, in fruit 8 –12 mm
long; ovary 1–1.5 mm long, style 2 – 5 mm long, stigmas 8 – 20 mm long; bracts peltate,
0.5 –1 mm diam., ciliolate. Fruit ellipsoid, 1–1.3 cm long, dehiscent, whitish; endocarp
body 0.6 – 0.8 cm long, smooth. — Fig. 10.
Distribution — Sumatra (also Riouw Archipelago), Malay Peninsula (Penang, Perak,
Trengganu to Singapore), Borneo, Celebes.
Habitat — Lowland forest, common in secondary forest.
Fig. 10. Streblus elongatus (Miq.) Corner. a. Part of inflorescence and in

the upper part two open pistillate flowers (one at anthesis, the other shortly
after) in section showing the bracts concealing the very young and, as yet,
minute staminate flowers, and in the upper part the abaxial sterile groove;
b, c. staminate flowers: 1 = pistillode; d, e. bracts (material from Singapore
Botanic Gardens).
e 1 mm 5 mm
b 3 mm
c
Uses — The species yields very hard, durable, strong, and heavy timber.
Notes — 1. Inflorescences are often deformed by glomerules of globose ‘bracteate’
structures (galls?).
2. The endocarp body is ejected for several meters. The exocarp is eaten by monkeys,
squirrels, and birds.
Section Taxotrophis
Streblus Lour. sect. Taxotrophis (Blume) Corner, Gard. Bull. Singapore 19 (1962) 218; Phytomor
phology 25 (1975) 5. — Taxotrophis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 77.
Diplocos Bureau in A.DC., Prodr. 17 (1873) 215.
Phyllochlamys Bureau in A.DC., Prodr. 17 (1873) 217. — Streblus Lour. sect. Phyllochlamys (Bureau)
Corner, Gard. Bull. Singapore 19 (1962) 217; Phytomorphology 25 (1975) 6.
Pseudotrophis Warb., Bot. Jahrb. Syst. 13 (1890) 291. — Streblus Lour. sect. Pseudotrophis (Warb.)
Corner, Gard. Bull. Singapore 19 (1962) 217; Phytomorphology 25 (1975) 5.
Dimerocarpus Gagnep., Bull. Mus. Hist. Nat. (Paris) 27 (1921) 441.
Trees or shrubs, mostly (or sometimes) with lateral or terminal thorns, dioecious.
Leaves distichous; lamina elliptic to narrowly elliptic, coriaceous, the margin often
spinulose-dentate and/or the acumen spinulose; petiole short; stipules free, lateral.
Inflorescences unisexual, the bracts basally attached; staminate inflorescences spicate
to subcapitate; pistillate inflorescences racemose to spicate to uniflorous; staminate
flowers 4-merous, tepals valvate; fruiting perianth more or less enlarged, not reflexed
in fruit. Drupe dehiscent or not.
Distribution — This section comprises six species and ranges from Sri Lanka to New
Guinea.
Morphology — All species bear thorns and/or have spinose-dentate leaf margins.
The thorns are formed terminally on leafy twigs or are lateral. In some species thorns
appear to be present on all specimens, in others they may be absent on individuals, or
regionally, or very rare (S. macrophyllus and S. perakensis). It is difficult to tell from
herbarium material whether or not the drupes are dehiscent.
This section is also distinct by the indumentum, occurring on the twigs unilaterally
only or unilateral more densely and on the petioles mostly adaxially only or adaxially
more densely.
KEY TO THE SPECIES
1a. Midrib of the lamina prominent above and acumen of the lamina spinulose-dentate,
with 1 or 3 teeth; lamina smooth and the margin often spinulose-dentate. — Wide-
spread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. S. ilicifolius
b. Midrib of the lamina impressed above or ± prominent only in the upper part of the
lamina and then the lamina usually scabrous; acumen of the lamina not spinulose-
dentate or, if apiculate, then in between 2 non-spinulose lobes . . . . . . . . . . . . . . . 2
2a. Plants usually with thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
b. Plants occasionally with thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3a. Thorns lateral; acumen of the lamina apiculate and often 2-lobed. — Sumatra, Java,
Lesser Sunda Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. S. spinosus
b. Thorns terminating leafy twigs; acumen of the lamina not apiculate and/or lobed . 4
4a. Plants usually with short-shoots bearing tufts of leaves and/or inflorescences; peti-
ole 0.1– 0.4(– 0.6) cm long, usually hairy adaxially; acumen of the lamina mostly
acute; leafy twigs usually distinctly hairy; tepals of the pistillate flower narrowly
ovate, acute. — Malay Peninsula, Java, Philippines, Celebes, Lesser Sunda Islands
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. S. taxoides
b. Plants rarely with short-shoots; petiole (0.2 –)0.4 – 0.7 cm long, often glabrous;
acumen of the lamina mostly obtuse; leafy twigs often subglabrous; tepals of the
pistillate flower broadly ovate to suborbicular, obtuse to rounded. — Malay Penin-
sula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. S. perakensis
5a. Staminate inflorescences spicate and 1.5 –14 cm long; pistillate inflorescences with
(1–)2 –10 flowers. — Malay Peninsula, Borneo, Philippines, Celebes . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. S. macrophyllus
b. Staminate inflorescences capitate or spicate and up to 0.8 cm long; pistillate inflo-
rescences with 1 (or 2) flowers. — Malay Peninsula . . . . . . . . . . 9. S. perakensis
7. Streblus ilicifolius (S. Vidal) Corner

Streblus ilicifolius (S. Vidal) Corner, Gard. Bull. Singapore 19 (1962) 227; Phytomorphology 25 (1975)
5, t. 2, 3F; Keng, Malaysian Seed Pl. (1969) f. 83. — Taxotrophis ilicifolia S. Vidal, Revis. Pl. Vasc.
Filip. (1886) 249; Merr., Interpr. Herb. Amboin. (1917) 188; Ridl., Fl. Malay Penins. 3 (1924) 322;
Corner, Gard. Bull. Singapore 10 (1939) 288; Wayside Trees Malaya ed.1 (1940) 693; Go, Tree Fl.
Sabah & Sarawak 3 (2000) 333.
Pseudotrophis laxiflora Warb., Bot. Jahrb. Syst. 13 (1891) 294.
Taxotrophis obtusa Elmer, Leafl. Philipp. Bot. 5 (1913) 1813.
Taxotrophis laxiflora Hutch., Bull. Misc. Inform. Kew (1918) 151. — Streblus laxiflorus (Hutch.)
Taxotrophis triapiculata Gamble, Bull. Misc. Inform. Kew (1913) 188.
Taxotrophis eberhardtii Gagnep., Fl. Indo-Chine 5 (1928) 700; Corner, Gard. Bull. Singapore 19
(1962) 229.
Taxotrophis macrophylla auct. non Boerl.: Burkill, Dict. Econ. Prod. Malay Penins. (1935) 2126.
Shrub or tree up to 10(– 20) m tall, much-branched, often with up to 4.5 cm long
lateral (or terminal) thorns, dioecious. Leafy twigs 1– 3 mm thick, brownish to whitish
puberulous, usually unilaterally only or more densely (or glabrous). Leaves distichous;
lamina elliptic to suborbicular to ovate or to obovate, 4 – 25 by 2 –10.5 cm, coriaceous,
apex acuminate to acute, acumen spinulose, usually with 3 teeth, base rounded to
cuneate, margin spinulose-dentate or entire; upper and lower surface glabrous; midrib
prominent above, lateral veins 8 –12 pairs, tertiary venation largely parallel to the lat-
eral veins; petiole 0.2 –1 cm long, whitish puberulous adaxially (hairy round about or
glabrous); stipules 0.3 – 0.8 cm long, free or basally connate, (sub)glabrous, caducous;
terminal bud aculeate in dry material. Staminate inflorescences axillary, solitary (or
in pairs), spicate; peduncle c. 0.1 cm long, sparsely minutely puberulous; spike 1– 5
cm long; flowers numerous; perianth 1–1.2 mm long, minutely puberulous; stamens
2 – 2.5 mm long, anthers 0.6 – 0.8 mm long; bracts basally attached, 0.3 –1.3 mm long,
sparsely minutely puberulous. Pistillate inflorescences axillary, solitary, (sub)racemose
or uniflorous; peduncle 0.3 –1.5 cm long, minutely puberulous; flowers 1– 3(–15);
pedicel 2 – 8 mm long, minutely puberulous; tepals 1– 2 mm long, ciliolate, in fruit the
outer ones up to 3 mm long, the inner ones up to 10 mm long; ovary 1–1.5 mm long,
style 0.5 –1 mm long, stigmas 2 – 3.5 mm long; bracts basally attached, 0.3 – 0.5 mm
long, ciliolate. Drupe 1–1.3 cm long, dehiscent, whitish; endocarp body subglobose to
ellipsoid, 0.6 – 0.8 cm long. — Map 4.
Distribution — Bangladesh, Myanmar, South China (incl. Hainan), Indochina, Thai-
land; in Malesia: Sumatra (northern), Malay Peninsula, Borneo, Philippines (Palawan,
Luzon, Mindoro, Basilan, Panay, Sulu Islands), Celebes (also Buton Island), Lesser
Sunda Islands (? Timor), Moluccas (Halmahera, Ambon, Ceram, Sula Islands, Key
Islands), New Guinea (Batanta Island).
Map 4. Distribution of Streblus ilicifolius (S. Vidal) Corner (broken line), S. macrophyllus Blume
(dot-dash line), and S. spinosus (Blume) Corner (dotted line).
Habitat — Dry rocky forest, especially in limestone hills and rocky coasts; at alti-
tudes up to c. 1100 m.
Uses — The timber is valuable.
Notes — 1. On the continent, in the Malay Peninsula, Sumatra and Borneo, the plants
often bear thorns, the margin of the lamina is usually spinulose dentate, and the pistillate
inflorescences contain usually 1– 3 flowers, whereas in the Philippines, Celebes, the
Moluccas, and New Guinea, the lamina is often entire and only the acumen spinulose.
Moreover, the plants are usually without thorns and the pistillate inflorescences often
bear more flowers, often up to c. 6, less commonly up to c. 15.
2. Terminal thorns are rare in Malesia; they are mainly found in Thailand.
3. The presence of this species in Timor could not be verified at L.
8. Streblus macrophyllus Blume

Streblus macrophyllus Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 80; Corner, Gard. Bull. Singa-
pore 19 (1962) 227; Phytomorphology 25 (1975) 5, t. 2, 3G; Go, Tree Fl. Sabah & Sarawak 3 (2000)
333. — Diplocos ? macrophyllus (Blume) Bureau in A.DC., Prodr. 17 (1873) 216. — Taxotrophis
macrophylla (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 359; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 38, excl. Taxotrophis ilicifolia; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 2126.
Pseudotrophis mindanaensis Warb. in Perkins, Fragm. Fl. Philipp. 1 (1905) 165; Elmer, Leafl. Philipp.
Bot. 5 (1913) 1815, ‘Taxatrophis mindanaensis’ in nota.
Paratrophis caudata Merr., Philipp. J. Sci., 1, Suppl. (1906) 183.
Taxotrophis balansae Hutch., Bull. Misc. Inform. Kew (1918) 151.
Dimerocarpus brenieri Gagnep., Bull. Mus. Hist. Nat. (Paris) 27 (1921) 441; Fl. Indo-Chine 5 (1928)
704, f. 84.
Tree up to 12 m tall, rarely with up to 0.5 cm long axillary thorns, dioecious or some-
times monoecious. Leafy twigs 1– 2.5 mm thick, minutely brownish puberulous, mostly
only unilaterally, or glabrous, young parts often ± compressed (often drying yellow-
ish). Leaves distichous; lamina elliptic to subovate to subobovate or to narrowly ovate,
6 – 22 by 2 –7.5 cm, often ± asymmetric, subcoriaceous, apex acuminate to subcaudate,
acumen acute, base rounded to obtuse to cuneate, margin subentire to crenate-dentate
to denticulate; upper surface glabrous; lower surface glabrous or whitish (minutely)
appressedly puberulous on the midrib; midrib ± impressed above, lateral veins 6 –10
pairs, tertiary venation subscalariform to reticulate; petiole 0.2 – 0.8 cm long, glabrous
or whitish minutely puberulous, often adaxially more densely; stipules 0.2 –1 cm long,
subglabrous or appressedly puberulous, subpersistent or caducous, terminal bud slen-
der, ± aculeate when dry. Staminate inflorescences axillary, solitary, spicate; pedun-
cle 0.2 – 0.4 cm long, minutely puberulous; spike 1.5 –14 cm long; flowers numerous;
perianth 1–1.2 mm long, minutely puberulous; stamens c. 2 mm long, anthers 0.6 – 0.8
mm long; bracts basally attached, 0.3 – 0.8 mm long, sparsely minutely puberulous.
Pistillate inflorescences axillary, solitary, racemose (or uniflorous); peduncle 0.2 –1
cm long, minutely puberulous; flowers (1–)2 –10; pedicel 0.5 –10 mm long, minutely
puberulous; tepals 1– 2 mm long, minutely puberulous, in fruit the outer ones 2 – 4 mm
long, the inner ones 7–10 mm long; ovary c. 1 mm long, style c. 1 mm long, stigmas
f c
g
5 mm
1 cm
4 mm
h
e
2 cm
1 cm 2.5 cm
a d b
Fig. 11. Streblus macrophyllus Blume. a. Leaf to show the intercostal veins in Bornean material;
b. fruiting spike; c. pistillate flower in section; d. dehiscing fruit; e, f. embryos in transverse and longi-
tudinal section: 1 = recurved lobe of outer cotyledon, 2 = inner cotyledon, 3 = outer cotyledon; g, h.
staminate flowers: 1 = pistillode (a – c: Jaheri 1234; d: Endert 3068; e: Santos 4215; g, h: Bur. Sci.
15485).
1.5 – 3 mm long; bracts basally attached, 0.3 –1 mm long, minutely puberulous. Drupe
1–1.4 cm long, dehiscent, whitish; endocarp body 0.8 –1 cm long. — Fig. 11; Map 4.
Distribution — Thailand, Indochina; in Malesia: Malay Peninsula, Borneo, Philip-
pines (Mindanao, Leyte, Basilan, Cebu, Surigao, Bohol), Celebes.
Habitat – Lowland forest.
9. Streblus perakensis Corner

Streblus perakensis Corner, Gard. Bull. Singapore 19 (1962) 223, t. 11; Phytomorphology 25 (1975) 3.
Shrub or tree up to 10 m tall, without or with lateral or terminal up to 1 cm long, often

± curved thorns, dioecious; often concentrations of persistent stipules (or scale-leaves)
and lenticels at branching points. Leafy twigs 1.5 – 3 mm thick, glabrous or minutely
whitish puberulous, unilaterally only or more densely. Leaves distichous; lamina nar-
rowly elliptic to subobovate, (2.5 –)5 –12 (–16) by (1–)2 – 5(–7) cm, often ± asymmetric,
subcoriaceous to coriaceous, apex acuminate, acumen obtuse to subacute, base rounded
to cordulate, margin irregularly dentate to denticulate or subentire; upper surface gla-
brous; lower surface glabrous or sparsely puberulous on the midrib; midrib ± impressed
above, lateral veins 6 – 8 pairs, tertiary venation reticulate; petiole (0.2 –)0.4 – 0.7 cm
long, glabrous or minutely whitish puberulous adaxially; stipules 0.1– 0.3 cm long,
appressedly puberulous, (sub)persistent. Staminate inflorescences axillary, solitary or
in pairs, spicate (or racemose), subsessile; spike (or raceme) 0.3 – 0.8 cm long; flowers
5 – 20, sessile (or up to 1 mm long pedicellate); perianth c. 1.5 mm long, minutely pu-
berulous; stamens c. 3 mm long, anthers c. 0.8 mm long; bracts basally attached, few,
0.2 – 0.5 mm long, sparsely puberulous. Pistillate inflorescences axillary, solitary or
in pairs, uniflorous, sessile or pedunculate; peduncle up to 0.4 cm long, subglabrous;
tepals 2 – 3 mm long, minutely puberulous, in fruit the outer ones 3 – 4 mm long, the
inner ones 10 –12 mm long, suborbicular; ovary c. 1.5 mm long, style c. 1 mm long,
stigmas 3 – 5 mm long; bracts 2, basally attached, 0.3 –1 mm long, minutely puberulous
or subglabrous. Drupe subglobose, 1–1.2 cm long, dehiscent, whitish; endocarp body
subglobose, c. 0.7 cm long.
Distribution — Peninsular Thailand; in Malesia: Malay Peninsula.
Habitat — Primary (and secondary) forest, sometimes on limestone hills; at altitudes
up to c. 500.
Notes — 1. In the very short and almost sessile staminate inflorescences this species
matches S. taxoides, from which it can be distinguished by somewhat longer petioles
and the absence of short-shoots bearing tufts of leaves and/or inflorescences. Also, the
shape of the tepals of the pistillate flower is distinctly different. The pistillate inflores-
cences are often sessile at anthesis. Spines are formed, but apparently not as frequently
and consistently as in S. taxoides. Concentrations of persistent scales and lenticels can
often be found on branching points.
2. Corner (1962) included S. perakensis in sect. Paratrophis. Corner (1975) main-
tained the species in this section with some doubt. However, the enlarged tepals in fruit
and the presence of thorns clearly indicate its correct position.
3. The distribution range of this species is quite small compared to the other species
of the section.
10. Streblus spinosus (Blume) Corner

Streblus spinosus (Blume) Corner, Gard. Bull. Singapore 19 (1962) 229; Backer & Bakh.f., Fl. Java
2 (1965) 16.
Urtica spinosa Blume, Bijdr. (1825) 507. — Taxotrophis spinosa (Blume) Steenis in Backer & Bakh.f.,
Fl. Java 2 (1965) 16.
Taxotrophis javanica Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 77, t. 26; Bureau in A.DC.,
Prodr. 17 (1873) 217; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 4, 6; Koord., Exkurs.-
Fl. Java 2 (1912) 88, f. 26; Hutch., Bull. Misc. Inform. Kew (1918) 149; Rendle, J. Bot. 63, Suppl.
(1925) 105.
Shrub or treelet up to 4 m tall, much-branched, with up to 1.5 cm long lateral thorns,

dioecious; short-shoots often present, bearing (clusters of) leaves and/or inflorescences.
Leafy twigs 1– 2.5 mm thick, whitish to brownish puberulous unilaterally or all over
(or glabrous). Leaves distichous; lamina elliptic to (sub)ovate or to narrowly ovate,
3 –12(–18) by 2 – 6(–7.5) cm, coriaceous, apex acuminate to subacute, acumen apicu-
late and usually 2-lobed, base rounded to obtuse, margin subspinulose-denticulate in
the upper part of the lamina or subentire; both surfaces glabrous; midrib ± impressed
above, lateral veins 6 –10 pairs, tertiary venation reticulate; petiole 0.2 – 0.5 cm long,
whitish puberulous unilaterally only or all over (or glabrous); stipules 0.2 – 0.3 cm
long, (sub)glabrous, caducous (or subpersistent); terminal bud spinose in dry mate-
rial. Staminate inflorescences axillary, solitary or in pairs, racemose; peduncle 0.1– 0.3
cm long, sparsely minutely puberulous; raceme 0.5 – 2.5 cm long; flowers numerous;
pedicel c. 0.1 mm long, minutely puberulous; perianth c. 1 mm long, minutely puberu-
lous; stamens 2 – 2.5 mm long, anthers c. 0.8 mm long; bracts basally attached, few,
0.5 –1.5 mm long, sparsely minutely puberulous. Pistillate inflorescences axillary, soli-
tary, uniflorous; peduncle 0.2 – 2 cm long, minutely puberulous; tepals 1–1.5 mm long,
minutely puberulous, in fruit hardly enlarged, up to 3 mm long; ovary c. 1 mm long,
style 0 – 4 mm long, stigmas 1– 2 mm long; bracts basally attached, few, 0.3 –1 mm
long, minutely puberulous. Drupe subglobose to ellipsoid, 0.6 –1 cm long, dehiscent (?),
whitish. — Map 4.
Distribution — Sumatra (also Enggano), Java, Lesser Sunda Islands (Bali, Flores,
Timor).
Habitat — Lowland and submontane forest at altitudes up to 1200 m; chiefly in open
country or rocky forest, on coral.
11. Streblus taxoides (B. Heyne ex Roth) Kurz

Streblus taxoides (B. Heyne ex Roth) Kurz, Forest Fl. Burma 2 (1877) 465; Corner, Gard. Bull. Singa-
pore 19 (1962) 225; Backer & Bakh.f., Fl. Java 2 (1965) 16; Corner, Phytomorphology 25 (1975)
3, t. 1C, 2, 3A; Fl. Ceylon 1.2 (1977) 158, t. 29. — Trophis taxoides B. Heyne ex Roth, Nov. Pl.
Sp. (1821) 368. — Trophis taxiformis Spreng., Syst. Veg. 3 (1826) 902, nom. nov. illeg. — Phyllo
chlamys taxoides (B. Heyne ex Roth) Koord., Exkurs.-Fl. Java 2 (1912) 89; Merr., Philipp. J. Sci.
14 (1920) 247; Enum. Philipp. Flow. Pl. 2 (1923) 38; Koord., Atlas Baumart. Java (1924) t. 748;
690.
Trophis spinosa Roxb., Fl. Ind., ed. Carey 3 (1832) 762, non Willd. 1806, nec Blume 1826. — Epi
carpurus spinosus (Roxb.) Wight, Icon. Pl. Ind. Orient. 6 (1853) 7, t. 1962, p.p. — Phyllochlamys
spinosa (Roxb.) Bureau in A.DC., Prodr. 17 (1873) 218; Hook.f., Fl. Brit. India 5 (1888) 488;
Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 10; Ridl., Fl. Malay Penins. 3 (1924) 323.
Epicarpurus timorensis Decne., Nouv. Ann. Mus. Hist. Nat. 3 (1834) 499, t. 21; Span., Linnaea 15
(1841) 344.
Epicarpurus zeylanicus Thwaites, Hooker’s J. Bot. Kew Gard. Misc. 3 (1851) t. 11; 4 (1852) 1. — Stre
blus zeylanicus (Thwaites) Kurz, Forest Fl. Burma 2 (1877) 464; Corner, Gard. Bull. Singapore
19 (1962) 229; Phytomorphology 25 (1975) 5, t. 1A; Fl. Ceylon 1.2 (1977) 160, t. 30. — Diplocos
zeylanica (Thwaites) Bureau in A.DC., Prodr. 17 (1873) 215.
Taxotrophis roxburghii Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 78; Miq., Fl. Ned. Ind. 1, 2
(1859) 279.
Taxotrophis zeylanica Thwaites, Enum. Pl. Zeyl. (1864) 264.
Streblus microphyllus Kurz, Prelim. Rep. Forest Pegu (1875) App. A, cxviii; App. B, 84; Forest Fl.
Burma 2 (1877) 464, in clavi. — Streblus taxoides (B. Heyne ex Roth) Kurz var. microphylla (Kurz)
Kurz, Forest Fl. Burma 2 (1877) 465.
Phyllochlamys wallichii King ex Hook.f., Fl. Brit. India 5 (1888) 489; Ridl., Fl. Malay Penins. 3 (1924)
322; Corner, Wayside Trees Malaya ed.1 (1940) 690.
Taxotrophis caudata Hutch., Bull. Misc. Inform. Kew (1918) 149.
Phyllochlamys taxoides (B. Heyne ex Roth) Koord. var. parvifolia Merr., Enum. Philipp. Flow. Pl. 2
(1923) 38.
Taxotrophis poilanei Gagnep., Fl. Indo-Chine 5 (1928) 701.
Taxotrophis crenata Gagnep., Fl. Indo-chine 5 (1928) 702, t. 82. — Streblus crenatus (Gagnep.) Corner,
Gard. Bull. Singapore 19 (1962) 226.
Phyllochlamys tridentata Gagnep., Fl. Indo-Chine 5 (1928) 714.
Shrub or treelet up to 5 m tall, much-branched, with thorns up to 1.5 cm long, mostly

terminating (short) leafy twigs, dioecious; short-shoots often present, bearing (clusters
of) leaves and /or inflorescences. Leafy twigs 1– 2.5 mm thick, brown to whitish puberu-
lous unilaterally only or all over (or glabrous). Leaves distichous; lamina elliptic to nar-
rowly ovate or (sub)obovate, (1–)2 –10(–18) by (0.5–)1– 4.5(– 6.5) cm, subcoriaceous
to coriaceous, apex acuminate to acute (to rounded), acumen acute to obtuse, base
obtuse to rounded to cordulate, margin dentate to denticulate (to crenate), mainly in the
upper part of the lamina; both surfaces glabrous and smooth; midrib impressed above,
lateral veins 6 –12 pairs, tertiary venation reticulate; petiole 0.1– 0.4(– 0.6) cm long,
brown to whitish puberulous, adaxially only (or more all over); stipules 0.2 – 0.5 cm
long, ciliolate, subpersistent. Staminate inflorescences axillary, 1– 4 on short-shoots,
short-spicate to (sub)capitate (or to racemose), (sub)sessile; spike (or raceme) 0.4 – 0.8
cm long; flowers 8 –14, (sub)sessile (or up to 1 mm long pedicellate); perianth c. 1.5 mm
long, subglabrous or minutely puberulous; stamens 2 – 2.5 mm long, anthers 0.6 – 0.8
mm long; bracts basally attached, 0.2 – 0.6 mm long, minutely puberulous. Pistillate
inflorescences axillary, solitary (sometimes clustered on short-shoots), uniflorous; pe-
duncle 0.3 – 0.6 cm long, subglabrous; tepals ovate, 2 – 5 mm long, glabrous, in fruit up
to 2 cm long; ovary c. 1.5 mm long, style c. 1 mm long, stigmas 0.5 – 3 mm long; bracts
usually 2, ovate to narrowly ovate, c. 1.5 mm long, subglabrous. Drupe ellipsoid, 0.5 –1
cm long, dehiscent (?), whitish. — Fig. 12.
Distribution — Sri Lanka, India, Bhutan, Myanmar, South China (Hainan), Thailand,
Indochina; in Malesia: Malay Peninsula, Java, Philippines (Mindoro, Palawan), Celebes
(south-eastern), Lesser Sunda Islands (Flores, Timor).
Habitat — Lowland forest in rocky or dry places.
1 cm
1 cm
1 cm
1 mm
e a b c d
Fig. 12. Streblus taxoides (B. Heyne ex Roth) Kurz. a. Ripe fruit in section: 1 = cotyledon, 2 = scar of
the cotyledon cut off, 3 = endocarp, 4 = exocarp; b. embryo in section; c. embryo from the lower side;
d. staminate inflorescence in section; e. pistillode (a – c: material from Singapore Botanic Gardens;
d, e: Rabil 380, Thailand).
Notes — 1. This species can be easily confused with S. spinosus. In S. taxoides, the
thorns are normally borne at the end of leafy twigs, the acumen of the lamina has an
acute apex, the stipules do not form an aculeate terminal bud and are subpersistent. In
the other species the thorns are borne laterally on twigs, the acumen of the lamina is
normally 2-lobed with a small tooth in between these lobes, the stipules form an aculeate
terminal bud and are caducous.
2. Staminate inflorescences are or can be subinvolucrate, due to enlargement of the
bracts, at least in Sri Lanka, India, and Thailand. In the Malesian region the bracts are
(always?) small.
3. In Thailand the tepals of the pistillate flowers are sometimes narrowly ovate.
4. Streblus zeylanicus is included as differentiating characters are absent.
TROPHIS
Trophis P. Browne, Civ. Nat. Hist. Jamaica (1756) 357, nom. cons.; Trécul, Ann. Sci. Nat., Bot. sér.
3, 8 (1847) 146; Bureau in A.DC., Prodr. 17 (1873) 251; Baill., Hist. Pl. 6 (1875) 192; Benth. &
Hook.f., Gen. Pl. 3 (1880) 365; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 73; Bot.
Jahrb. Syst. 40 (1908) 543; W.C. Burger, Ann. Missouri Bot. Gard. 49 (1962) 6; Corner, Gard.
Bull. Singapore 19 (1962) 230; C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 352; Fl.
Neotrop. Monogr. 83 (2001) 33.
Bucephalon L., Sp. Pl. (1753) 1190.
Olmedia Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1 (1798) 257; C.C. Berg, Fl. Neotrop. Monogr. 7
(1972) 14. — Trophis P. Browne sect. Olmedia (Ruiz & Pav.) C.C. Berg, Proc. Kon. Ned. Akad.
Wetensch. C, 91 (1988) 354.
Malaisia Blanco, Fl. Filip. (1837) 789; ed. 2 (1845) 543; ed. 3, 3 (1879) 196; Planch., Ann. Sci. Nat.,
Bot. sér. 4, 3 (1855) 293; Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 75; Miq., Fl. Ned. Ind.
1, 2 (1859) 281; Bureau in A.DC., Prodr. 17 (1873) 221; Benth. & Hook.f., Gen. Pl. 3 (1880) 360;
Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 76; Boerl., Handl. Fl. Ned. Ind. 3 (1900)
315; Corner, Gard. Bull. Singapore 19 (1962) 240. — Trophis P. Browne sect. Malaisia (Blanco)
C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 354.
Dumartroya Gaudich., Voy. Bonite, Bot. (1844) 165, t. 97.
Cephalotrophis Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 75.
Maillardia Frapp. ex Duch., Notes sur l’île de la Réunion, Annexe Phanerogamique (1862) 148; Bureau
in A.DC., Prodr. 17 (1873) 220; Leandri, Fl. Madagasc. fam. 55 (1952) 15. — Trophis P. Browne
sect. Maillardia (Frapp. ex Duch.) Corner, Gard. Bull. Singapore 19 (1962) 230.
Calpidochlamys Diels, Bot. Jahrb. Syst. 67 (1935) 172. — Trophis P. Browne sect. Calpidochlamys
(Diels) Corner, Gard. Bull. Singapore 19 (1962) 230.
Skutchia Pax & K. Hoffm. ex C.V. Morton, J. Wash. Acad. Sci. 27 (1937) 306.
Caturus Lour. auct. non L. (1767): Fl. Cochinch. (1790) 612; Baill., Hist. Pl. 6 (1875) 193.
Trees or shrubs (scandent in T. scandens), dioecious, unarmed, uncinate hairs lacking

or present. Leaves distichous; lamina pinnately veined; stipules free, lateral. Inflores
cences unisexual, racemose, spicate, subcapitate, or discoid-capitate and involucrate,
bracteate. Staminate inflorescences multi- to pluriflorous, flowers with 3 or 4 tepals,
valvate or imbricate in bud; stamens 3 or 4, inflexed in bud. Pistillate inflorescences
multi- to uniflorous; flowers with 4 connate tepals, equal in size and forming a tubular
or a collar-shaped perianth; ovary free or adnate to the perianth; stigmas 2, equal. Fruit
ing perianth enlarged, fleshy, orange or red, enclosing the free and indehiscent or adnate
fruit, endocarp crustaceous to woody; seed without endosperm, embryo with thick, flat,
equal or (very) unequal cotyledons, radicle short or rather long.
Distribution — The genus comprises nine species in the Old World (not on the Afri
can continent) and New World.
Subdivision — The genus can be subdivided into six sections (Berg 1988): Calpido
chlamys (one species in Malesia), Echinocarpa (one species, T. involucrata W.C. Burger,
in Central America), Maillardia (two species in Madagascar and Réunion), Malaisia
(one species in SE Asia), Olmedia (one species, T. caucana (Pittier) C.C. Berg, in the
Neotropics, and Trophis (3 species in the Neotropics, see Berg 2001). This subdivision
replaces that of Corner (1962).
Morphology — This genus shows pronounced morphological similarities to Streblus
but differs in the connate tepals of the pistillate flower.
Akad. Wetensch. C, 91 (1988) 345 – 362. — Berg, C.C., Fl. Neotrop. Monogr. 83 (2001) 32. — Corner,
E.J.H., The classification of Moraceae. Gard. Bull. Singapore 19 (1962) 187– 252.
KEY TO THE SPECIES
1a. Tree; staminate inflorescences spicate, 6 – 26 cm long; pistillate inflorescences

1.5 – 4 cm long. — Eastern Malesia . . . . . . . . . . . . . . . . . . . . 1. T. philippinensis
b. Climber; staminate inflorescences spicate to subcapitate, 0.5 – 3.5(– 5) cm long;

pistillate inflorescences capitate to short-spicate, 0.3 –1.5 cm long. — Widespread
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. T. scandens
Section Calpidochlamys
Trophis P. Browne sect. Calpidochlamys (Diels) Corner, Gard. Bull. Singapore 19 (1962) 230; C.C.
Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988) 356. — Calpidochlamys Diels, Bot. Jahrb.
Syst. 67 (1935) 172.
Trees. Staminate inflorescences spicate; tepals 4, imbricate in the bud. Pistillate

inflorescences spicate; fruiting perianth enlarged, fleshy, dark purple to black. Fruit
adnate to the perianth; cotyledons subequal.
Distribution — This section comprises only T. philippinensis.
1. Trophis philippinensis (Bureau) Corner

Trophis philippinensis (Bureau) Corner, Gard. Bull. Singapore 19 (1962) 231. — Uromorus philip
pinensis Bureau in A.DC., Prodr. 17 (1873) 237; S. Vidal, Cat. Met. Pl. Leños. (1880) 43. — Para
trophis philippinensis (Bureau) Fern.-Vill., Nov. App. (1880) 98; S. Vidal, Phan. Cuming. Philipp.
(1885) 145; Revis. Pl. Vasc. Filip. (1886) 250; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 36.
Sloetia minahassae Koord., Versl. Minahasa (1898) 645; Suppl. Fl. Celebes 2 (1922) pl. 5; 3 (1922) 3.
Paratrophis grandifolia Elmer, Leafl. Philipp. Bot. 5 (1913) 1814; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 36.
Calpidochlamys branderhorstii Diels, Bot. Jahrb. Syst. 67 (1935) 173. — Trophis branderhorstii
(Diels) Corner, Gard. Bull. Singapore 19 (1962) 231.
Calpidochlamys drupacea Diels, Bot. Jahrb. Syst. 67 (1935) 173. — Trophis drupacea (Diels) Corner,
Gard. Bull. Singapore 19 (1962) 231.
Tree up to 30 m tall. Leafy twigs 1.5 – 2.5 m thick, brownish to whitish puberulous,
mostly drying dark brown with conspicuous lenticels. Leaves distichous; lamina nar-
rowly elliptic to elliptic to subobovate, (4 –)10 – 20(– 28) by (2 –)4 – 9(–14), usually
± asymmetric, coriaceous to subcoriaceous, apex acuminate, base cuneate to obtuse (to
almost rounded), margin entire, often ± revolute, at least towards the base; upper surface
glabrous; lower surface brownish to whitish puberulous to strigillose on the veins or
(sub)glabrous; midrib prominent above, lateral veins 9 –16(– 20) pairs, tertiary venation
scalariform; petiole 0.8 – 2.5 cm long, brownish to whitish puberulous to subglabrous;
stipules 0.5 –1 cm long, sparsely appressedly brownish to whitish puberulous, caducous.
Staminate inflorescences axillary, solitary or in pairs, spicate; peduncle 0.3 – 2 cm long,
(sub)glabrous; spike 6 – 26 cm long; staminate flowers numerous; perianth 1–1.2 mm
long, sparsely hairy to glabrous; stamens c. 2 mm long, anthers c. 0.5 mm long; bracts
peltate, 0.5 –1 mm diam., (sub)glabrous or sparsely ciliolate. Pistillate inflorescences
axillary, solitary or in pairs, spicate; peduncle 0.2 – 0.6 cm long, subglabrous; spike
1.5 – 4 cm long; flowers 2 –10; perianth c. 2 mm long, sparsely minutely puberulous,
4-dentate; ovary c. 1 mm long, style 1– 2 mm long, stigmas 2 – 3.5 mm long; bracts
peltate, 0.5 –1 mm diam., sparsely ciliolate. Fruiting perianth ellipsoid to subglobose,
when fresh up to 3 cm long, when dry 1.5 – 2 cm long, dark purple to black at maturity.
— Fig. 13.
5 mm
e f g
2 cm
1 cm
5 mm
1 mm
a c d b
Fig. 13. Trophis philippinensis (Bureau) Corner. a. Leafy twig with staminate inflorescence; b. pistillate
inflorescence at anthesis; c. pistillate flower in section; d. fruit in section: 1 = thickened hilar plug of
endocarp, 2 = endocarp, 3 = fruiting perianth; e – g. embryo (a: Ramos 1638; b, c: BW 363; d: Cel./V-
193; e – g: Branderhorst 273).
Distribution — Borneo (Sarawak), Philippines (Luzon, Samar, Mindanao, Palawan),

Celebes, Lesser Sunda Islands (Sumba, Flores), Moluccas (Halmahera), New Guinea
(also Yapen and New Britain).
Habitat — Primary (and secondary) forest at altitudes up to 1500 m.
Uses — Young leaves are cooked and eaten; wood is used for house building.
Section Malaisia
Trophis P. Browne sect. Malaisia (Blanco) C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C, 91 (1988)
354. — Malaisia Blanco, Fl. Filip. (1837) 789
Climbers. Staminate inflorescences spicate to subcapitate; tepals 3 (or 4), valvate in

the bud. Pistillate inflorescences (sub)capitate; perianth enlarged in fruit, fleshy, red.
Fruit free, somewhat drupaceous; cotyledons unequal.
Distribution — This section comprises only T. scandens.
2. Trophis scandens (Lour.) Hook. & Arn.

Trophis scandens (Lour.) Hook. & Arn., Bot. Beechey Voy. (1837) 214; C.C. Berg, Proc. Kon. Ned.
Akad. Wetensch. C, 91 (1988) 354. — Caturus scandens Lour., Fl. Cochinch. (1790) 612. — Malaisia
scandens (Lour.) Planch., Ann. Sci. Nat., Bot. sér. 4, 3 (1855) 293; Blume, Ann. Mus. Bot. Lugduno-
Batavi 2 (1856) 76; Merr., Philipp. J. Sci., 1, Suppl. (1906) 43; Koord., Exkurs.-Fl. Java 2 (1912) 86;
Merr., Sp. Blancoan. (1918) 122; Enum. Philipp. Flow. Pl. 2 (1923) 37; Comm. Fl. Cochinch. (1935)
132; Specht, Rec. Amer.-Austral. Sci. Exped. Arnhem Land, 3, Bot. Pl. Ecol. (1958) 218, 330; T.S.
Liu, Ill. Lign. Pl. Taiwan 2 (1962) 747; H.L. Li, Woody Fl. Taiwan (1963) 127, f. 38; Backer &
Bakh.f., Fl. Java 2 (1965) 15. — Alchornea scandens (Lour.) Müll.Arg., Linnaea 34 (1865) 170;
Müll.Arg. in A.DC., Prodr. 15, 2 (1866) 906. — Malaisia tortuosa Blanco var. scandens (Lour.)
Bureau in A.DC., Prodr. 17 (1873) 222.
Morus javanica Blume, Bijdr. (1825) 488. — Cephalotrophis javanica (Blume) Blume, Ann. Mus. Bot.
Lugduno-Batavi 2 (1856) 76, t. 27; Miq., Fl. Ned. Ind. 1, 2 (1859) 281.
Malaisia tortuosa Blanco, Fl. Filip. (1837) 789; Blume, Ann. Mus. Bot. Lugduno-Batavi 2 (1856) 75;
Miq., Fl. Ned. Ind. 1, 2 (1859) 282; Benth., Fl. Australia 6 (1873) 180; Bureau in A.DC., Prodr. 17
(1873) 221; Fern.-Vill., Nov. App. (1880) 198; S. Vidal, Sin. Gen. Pl. Leños Filip. (1883) 39, t. 86
f. B; Phan. Cuming. Philipp. (1885) 145; Revis. Pl. Vasc. Filip. (1886) 250; Merr., Publ. Gov. Lab.
Philipp. 27 (1905) 78; Ridl., Fl. Malay Penins. 3 (1924) 325; Gagnep., Fl. Indo-Chine 5 (1928) 696
(sub var. scandens), t. 83: 1– 4.
Dumartroya fagifolia Gaudich., Voy. Bonite, Bot. (1844) 165, t. 97.
Malaisia viridescens Planch., Ann. Sci. Nat., Bot. sér 4, 3 (1855) 293. — Malaisia tortuosa Blanco
var. viridescens (Planch.) Bureau in A.DC., Prodr. 17 (1873) 222.
Malaisia acuminata Planch., Ann. Sci. Nat., Bot. sér. 4, 3 (1855) 294. — Malaisia tortuosa Blanco
var. acuminata (Planch.) Bureau in A.DC., Prodr. 17 (1873) 222.
Malaisia cunninghamii Planch., Ann. Sci. Nat., Bot. sér. 4, 3 (1855) 294.
Cephalotrophis ? puberula Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 416. — Malaisia puberula (Miq.)
Bureau in A.DC., Prodr. 17 (1873) 222.
Caturus torulosus Seem., Fl. Vit. (1868) 254.
Malaisia tortuosa Blanco var. racemosa Bureau in A.DC., Prodr. 17 (1873) 222.
Malaisia scandens (Lour.) Planch. var. rolfei K. Schum., Bot. Jahrb. Syst. 9 (1888) 199.
Malaisia blancoi Elmer, Leafl. Philipp. Bot. 5 (1913) 1812.
Climber, or shrub with climbing branches, these with long internodes and rudimen-
tary leaves. Leafy twigs 1.5 – 2.5 mm thick, brownish to whitish puberulous, partly
with uncinate hairs, mostly drying dark brown with conspicuous lenticels. Leaves dis-
tichous; lamina narrowly elliptic to narrowly ovate, 4 –16(– 20) by 1.5 – 6.5(– 8), usu-
ally ± asymmetric, subcoriaceous, apex acuminate, base rounded to subcordate (or to
subcuneate), margin entire or denticulate to dentate towards the apex, often ± revolute,
at least towards the base; upper surface minutely whitish puberulous on (the lower part
of) the midrib; lower surface sparsely whitish puberulous on the (main) veins; midrib
prominent above, lateral veins 7–12 pairs, tertiary venation (sub)scalariform; petiole
0.3 –1(–1.5) cm long, (sparsely) puberulous; stipules 0.1– 0.5 cm long, sparsely whitish
appressedly puberulous, caducous or subpersistent. Staminate inflorescences axillary,
solitary, in pairs or up to 6 together on short-shoots, spicate; peduncle 0.5 –1.5 cm long,
puberulous to tomentellous; spike 0.5 – 3.5(– 5) cm long, often interrupted; perianth
c. 1.5 mm long, minutely puberulous; stamens c. 2.5 mm long, anthers c. 0.8 mm long;
bracts basally attached, ovate to narrowly ovate, 0.5 –1 mm long, minutely puberulous.
Pistillate inflorescences axillary, solitary, in pairs or up to 6 together on short-shoots,
capitate to short-spicate (or uniflorous); peduncle 0.4 – 2 cm long, puberulous to tomen-
tellous; head 0.3 – 0.5 cm diam., spike up to 1.5 cm long; flowers 1– 8; perianth 1–1.5
mm long, glabrous except for the densely white puberulous apex, faintly 4-dentate
to subentire; ovary c. 1 mm long, style 1– 2 mm long, stigmas 5 –15 mm long; bracts
basally attached, ovate to subulate, 0.5 –1 mm long, densely white puberulous to sub-
tomentellous. Fruiting perianth ellipsoid, when dry 0.6 – 0.8 cm long, dark pink to red
at maturity. — Fig. 14.
Distribution — South China (incl. Hainan and Taiwan), Indochina, Thailand, Aus-
tralia (Darwin, Arnhem Land, Queensland, New South Wales) and Pacific: New Cale
2 cm
1 mm
1 cm
a
5 mm
c
f
4 mm
d e
Fig. 14. Trophis scandens (Lour.) Hook. & Arn. a. Leafy twig with staminate inflorescences; b, c.
staminate flowers; d. pistillate inflorescence in section, just after anthesis; e. pistillate inflorescence in
fruit; f. embryo (a: Pullen 1167; b, c: PNH 15730; d – f: Corner s.n., New Guinea).
donia, Lord How Island, Vanuatu, Fiji; in Malesia: Sumatra, Malay Peninsula, Borneo,
Java, Philippines, Celebes, Lesser Sunda Islands (Bali, Sumbawa, Flores, Timor, Wetar),
Moluccas (Sula Islands, Banda, Ambon, Tanimbar Islands), New Guinea.
Habitat — Forest, mainly forest edges, floodplain forest, in humid to dry habitats,
sometimes on limestone at altitudes up to 1300 m.
Tribe Antiaropsideae
Trees, dioecious, without uncinate hairs. Leaves distichous or to laxly spirally arranged;
lamina entire; venation subscalariform to reticulate; stipules free, lateral to almost
fully amplexicaul. Inflorescences axillary, unisexual; receptacle discoid to urceolate,
involucrate with imbricate or few ± scattered basally attached bracts, possibly interfloral
bracts in Antiaropsis. Flowers free, tepals 4 or 5, free; stamens (2–)4 – 6, straight in the
bud; pistillode present; ovary free, stigmas filiform, 2 and equally or unequally long,
or 1. Fruit a dehiscent drupe.
Distribution — The tribe comprises two genera, Antiaropsis confined to New Guin-
ea, with 2 species and Sparattosyce, confined to New Caledonia, with one (or two?)
species.
Morphology — The tribe shows similarities to the Castilleae in the structure of the
inflorescences but it clearly differs in the absence of connate tepals, the non-fleshy
tepals of pistillate flowers in fruit, the free dehiscent drupes, and the absence of self-
pruning branches (as part of the architectural model of Cook (Hallé & Oldeman, Essai
sur l’architecture et la dynamique de croissance des arbres tropicaux (1970); Berg, Acta
Bot. Neerl. 26 (1977)).
Antiaropsis
Antiaropsis K. Schum., Fl. Kais. Wilh. Land (1889) 40; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 327;
K. Schum. & Lauterb., Fl. Schutzgeb. Südsee (1900) 267; Engl. in Engl. & Prantl, Nat. Pflanzen-
fam. Nachtr. 2 (1908) 96; Diels, Bot. Jahrb. Syst. 67 (1935) 174; Corner, Gard. Bull. Singapore 19
(1962) 249, ‘Antiaropsi’.
Trees or shrubs, dioecious. Leaves distichous; lamina pinnately veined; stipules free,
lateral. Inflorescences discoid and involucrate, with interfloral bracts (?), pedunculate.
Staminate flowers with 4 or 5 (or 6), (almost) free, imbricate tepals; stamens (2 –)
3 – 5(– 6). Pistillate flowers numerous, tepals 4, free; stigmas 2, filiform. Fruit a dehis-
cent drupe; seed without endosperm, cotyledons equal, rather thick, involute, radicle
apical, short.
Distribution — The genus comprises two closely related species and is only known
from New Guinea.
Morphology — The pistils are surrounded by 4, ± conduplicate tepals. In the pistil-
late inflorescences of A. decipiens, in addition to the tepals, similar, somewhat larger
and flat, structures occur on the receptacle, which could be regarded as interfloral bracts,
but might be ‘displaced’ tepals. Such ‘interfloral bracts’ have not yet been detected in
the material available of staminate inflorescences.
Chemistry — α-antiarin and antioside, cardenolides known from Antiaris have also
been found in seeds of Antiaropsis (Bisset, Ann. Bogor. 2 (1957) 219).
KEY TO THE SPECIES
1a. Lamina (usually) longer than 10 cm, often ± scabrous beneath; pistillate inflo
rescences with numerous flowers; involucral bracts (rather) sparsely hairy; up to
1250 m altitude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. A. decipiens
b. Lamina up to 10 cm long, smooth beneath; pistillate inflorescences with one flower;
involucral bracts densely hairy; 1400 – 3200 m altitude . . . . . . . . . . 2. A. uniflora
1. Antiaropsis decipiens K. Schum.

Antiaropsis decipiens K. Schum., Fl. Kais. Wilh. Land (1889) 40; K. Schum. & Lauterb., Fl. Schutz
geb. Südsee (1900) 267; Diels, Bot. Jahrb. Syst. 67 (1935) 174; Corner, Gard. Bull. Singapore. 19
(1962) 249.
Shrub or tree up to 12 m tall. Leafy twigs 1– 2.5 mm thick, appressedly puberulous.

Leaves distichous; lamina elliptic to subobovate or subovate or to narrowly ovate,
(2.5 –)6 –14(– 23) by (1.5 –)2.5 – 6(–10.5) cm, coriaceous to subcoriaceous, apex acu-
minate to subcaudate, base cuneate to obtuse (to truncate), margin entire (or coarsely
dentate towards the apex to lobate); upper surface (sub)glabrous; lower surface sparsely
appressedly puberulous on the main veins, ± scabrous or smooth; midrib prominent
above, lateral veins (5 –)7–14(–17) pairs, tertiary venation (sub)scalariform; petiole
0.5 –1.3 cm long, sparsely puberulous; stipules 0.2 – 0.6 cm long, puberulous, caducous.
Staminate inflorescences axillary, solitary; peduncle 8 –15 mm long, puberulous, mostly
with some small scattered bracts; head discoid, 5 –15 mm diam., involucral bracts in
3 or 4 rows, ovate, 1– 4 mm long, appressedly puberulous, orange to red; tepals 4 or 5
(or 6), 1.5 – 2 mm long, spathulate-cucullate, puberulous; stamens (2 –)3 – 5(– 6), 2 – 3
mm long, filaments free, 0.5 – 0.8 mm long, anthers 1–1.5 mm long, apiculate or not.
Pistillate inflorescences axillary, solitary, at maturity of the fruits pendulous; peduncle
1.2 – 3.5 cm long, puberulous, mostly with some small scattered bracts; head discoid,
1–1.5 cm, in fruit up to 3 cm diam.; involucral bracts in 5 or 6 rows, ovate to narrowly
ovate, 3 –10 mm long, appressedly puberulous, in fruit red; flowers numerous; tepals
narrowly elliptic to narrowly ovate, c. 5 mm, in fruit up to 10 mm long, ± conduplicate,
appressedly puberulous, in fruit orange to red; style 2 – 3 mm long, stigmas filiform,
1.5 – 3 mm long, equal or unequal in length; ‘interfloral bracts’ subovate to narrowly
elliptic 5 –7 mm long, in fruit up to 10 mm long, flat, appressedly puberulous. Endocarp
body ellipsoid, 6 –7 mm long, blackish, the exocarp white. — Fig. 15a – j.
Distribution — New Guinea.
Habitat — Evergreen forest at altitudes up to 1250 m.
Notes — 1. Shrubs as little as 0.5 m high may already flower.
2. Var. parvifolia Diels (1935), with lamina 4.5 – 6.5 by 2.5 cm and 4 or 5 pairs of
lateral veins, prominent beneath, is not included as the type could not be examined and
its identity is uncertain because of lack of details of the venation. For the same reasons
the type could not be included in the other species.
1 cm
5 mm
3 cm
c
i d
j 1 cm
1 cm
h e
1 cm
g f
10 cm
1 cm
k l m b a
3 mm
Fig. 15. a – j: Antiaropsis decipiens K. Schum. a. Leaf; b. sapling leaf; c. staminate inflorescence in
section; d. staminate flower: 1 = pistillode; e. infructescence in section; f. drupe in section: 1 = exocarp,
2 = endocarp, 3 = seed; g –j. embryo: 1 in g and i = radicle, 1 in j = cotyledons. — k – m: Antiaropsis
parvifolia C.C. Berg. k. Leaf; l. pistillate inflorescence in section; m. pistil in section (a: Nooroh 18l;
b, e – j: Janowsky 194; c, d: Schlechter 16045; k– m: McAdam s.n., Wau).
3. The blackish endocarp body, the white exocarps, and the red to orange coloured
tepals establish a set of contrasting colours, common in angiosperms to attract birds for
dispersal.
4. This species is pollinated by thrips, as has been described in detail by Zerega et
al., Int. J. Pl. Sci. 165 (2004) 1017–1026).
2. Antiaropsis uniflora C.C. Berg

Antiaropsis uniflora C.C. Berg, Blumea 50 (2005) 539.
?Antiaropsis decipiens K. Schum. var. parvifolia Diels, Bot. Jahrb. Syst. 67 (1935) 174; Corner, Gard.
Bull. Singapore 19 (1962) 250.
Tree up to 23 m tall. Leafy twigs 1– 2 mm thick, appressed puberulous. Leaves

distichous; lamina elliptic, 3 –10 by 1.5 – 3.5 cm, coriaceous to subcoriaceous, apex
acuminate to subcaudate, base (sub)cuneate, margin entire; upper surface glabrous;
lower surface sparsely appressed puberulous on the main veins, smooth; midrib promi-
nent above, lateral veins 8 –12 pairs, tertiary venation reticulate; petiole 0.4 – 0.8 cm
long, sparsely puberulous; stipules 0.2 – 0.4 cm long, puberulous, caducous. Staminate
inflorescences unknown. Pistillate inflorescences axillary, solitary; peduncle 1– 2.5
cm long, puberulous, without bracts; head discoid, c. 5 mm diam., in fruit up to c. 10
mm diam.; involucral bracts in 4 or 5 rows, semicircular to narrowly elliptic, 2 – 8 mm
long, densely white appressedly puberulous, the inner ones obtuse; flower one; tepals
4, narrowly elliptic to elliptic, c. 5 mm, in fruit up to 10 mm long, ± conduplicate, ap-
pressedly puberulous, in fruit orange to red; style 1.5 – 2 mm long, stigmas filiform,
1.5 – 2 mm long. Endocarp body ellipsoid, 7– 8 mm long. — Fig. 15k – m.
Habitat — Evergreen forest at altitudes between c. 1400 and 3200 m.
Notes — 1. Even though the material presently available is sparse, this taxon, re-
garded as a variety by Diels (1935) and by Corner (1962), is distinct enough to be
recognised as a species, as considered by Corner in the Flora Malesiana manuscript.
It is distinct from the lowland A. decipiens in the habit, becoming a tree of more than
20 m tall, the small leaves, up to 10 cm long, the lack of bracts on the peduncle of the
pistillate inflorescences, the broader and more densely hairy involucral bracts, and the
presence of a single flower in the pistillate inflorescence.
2. The same contrasting colours as found in the inflorescences of A. decipiens prob-
ably also occur in this species in fruit.
3. As the type of A. decipiens var. parvifolia is not extant, it is not certain whether it
belongs to this species or whether it is just a small-leaved form of A. decipiens.
Tribe Artocarpeae
Artocarpeae R.Br. in Tuckey, Narr. Exped. Zaire, App. 5 (1818) 454; Gaudich. in Freyc., Voy. Uranie,
Bot. (1830) 511; Corner, Gard. Bull. Singapore 19 (1962) 231; C.C. Berg, Proc. Kon. Ned. Akad.
Wetensch. C, 91 (1988) 357; Blumea 50 (2005) 535. — Euartocarpeae Trécul, Ann. Sci. Nat., Bot.
sér. 3, 8 (1847) 108.
Trees or shrubs (or climbers?), monoecious or dioecious, with or without uncinate

hairs. Leaves spirally arranged or distichous; stipules fully amplexicaul to lateral or
intrapetiolar, free or connate. Inflorescences usually unisexual, axillary or cauliflorous,
spicate to capitate and then clavate, globose, or discoid, pedunculate, bracteate; inter
floral bracts often peltate or subpeltate varying to spathulate, clavate, rod-shaped to
spine-shaped. Staminate flowers numerous, with 2 – 4 connate tepals, stamens 1– 4,
straight in the bud, pistillode rarely present. Pistillate flowers free or connate, tepals
2 – 4(– 5) and connate or absent (in Treculia), ovary free or adnate to the perianth, stig-
mas 1 or 2. Fruit large, free, with a dry pericarp or ± drupaceous, indehiscent embed-
ded in the pulpy part of the (many-seeded) infructescence; seed without or with scanty
endosperm, embryo curved or straight, with thick, equal or unequal cotyledons.
Distribution — This tribe comprises five genera of which four, Artocarpus (c. 45
spp.), Hullettia (2 spp.), Parartocarpus (2 spp.), and Prainea (2 spp.) occur in Asia
and are speciose in Malesia, two of them, Artocarpus and Parartocarpus extending to
Australia and /or the Pacific. The fifth genus, Treculia (with 3 spp.) occurs in continental
Africa and Madagascar (see Berg, Bull. Jard. Bot. Belg. 47 (1977) 378).
Morphology — The very large infructescences formed by some species are remark-
able and in Artocarpus resemble Durian fruits. Also notable are the wide range of shapes
derived from the basically peltate bracts and the presence of glands or glandular spots
in species of Artocarpus subg. Pseudojaca.
KEY TO THE GENERA
1a. Inflorescences involucrate with relatively large basally attached bracts at the base of
the globose receptacle, or bracts at the margin of the turbinate to discoid receptacle;
staminate flowers usually with 2 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Inflorescences without subtending bracts; staminate flowers with one stamen . . . 3
2a. Stipules free, linear, subpersistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hullettia
b. Stipules fused, broader, caducous . . . . . . . . . . . . . . . . . . . . . . . . . Parartocarpus
3a. Trees dioecious; pistillate flowers free . . . . . . . . . . . . . . . . . . . . . . . . . . . Prainea
b. Trees monoecious; pistillate flowers connate . . . . . . . . . . . . . . . . . . Artocarpus
ALTERNATIVE KEY TO THE GENERA
1a. Leaves spirally arranged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b. Leaves distichous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2a. Stipules fully amplexicaul (leaving an annular scar) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Artocarpus subg. Artocarpus
b. Stipules not fully amplexicaul . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Stipules lateral, subulate, subpersistent . . . . . . . . . . . . . . . . . . . . . . . . . . Hullettia
b. Stipules intrapetiolar, broader, caducous . . . . . . . . . . . . . . . . . . . . Parartocarpus
4a. Trees dioecious; pistillate flowers free . . . . . . . . . . . . . . . . . . . . . . . . . . . Prainea
b. Trees monoecious; pistillate flowers connate . . . . Artocarpus subg. Pseudojaca
ARTOCARPUS
Artocarpus J. & G. Forst., Char. Gen. Pl., ed. 2 (1776) 101, t. 51, 51a, nom. cons.; L.f., Suppl. (1781)
61, 411; Lam., Encycl. 3 (1789) 207; Willd., Sp. Pl. 4 (1805) 188; Pers., Syn. Pl. 2 (1807) 531;
Blume, Bijdr. (1825) 479; Roxb., Fl. Ind., ed. Carey 3 (1832) 521; Blanco, Fl. Filip. (1837) 666;
Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 109; Miq., Fl. Ned. Ind. 1, 2 (1859) 284; Fl. Ned. Ind.,
Eerste Bijv. (1861) 417; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211; Blanco, Fl. Filip. ed. 3, 3
(1879) 73; Benth. & Hook.f., Gen. Pl. 3 (1880) 376; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3,
1 (1888) 82; King in Hook.f., Fl. Brit. India 5 (1888) 539; King, Ann. Roy. Bot. Gard. (Calcutta) 2
(1889) 1; Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 2 (1900) 17; Boerl., Handl. Fl. Ned.
Ind. 3 (1900) 330, 370; Becc., For. Borneo (1902) 625; Koord. & Valeton, Bijdr. Boomsoort. Java
11 (1906) 11; Renner, Bot. Jahrb. Syst. 39 (1907) 363; Elmer, Leafl. Philipp. Bot. 2 (1909) 609;
Koord., Exkurs.-Fl. Java 2 (1912) 91; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 40; Ridl., Fl. Malay
Penins. 3 (1924) 351; Gagnep., Bull. Soc. Bot. France 73 (1926) 86; Fl. Indo-Chine 5 (1928) 731;
649; F.G. Browne, For. Trees Sar. & Brunei (1955) 349; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
113; Backer & Bakh.f., Fl. Java 2 (1965) 18; F.M. Jarrett, Blumea 22 (1975) 409; Kochummen,
Tree Fl. Malaya 3 (1978) 120; Corner, Wayside Trees Malaya ed. 3, 2 (1988) 511; Kochummen,
Tree Fl. Sabah & Sarawak 3 (2000) 187.
Sitodium Banks & Sol. ex Parkinson, J. Voy. South Seas (1773) 45; Thunb., Philos. Trans. 69 (1779)
465; Gaertn., Fruct. Sem. Pl. 1 (1788) 344, t. 71, 72; Fosberg, Amer. J. Bot. 26 (1939) 230; Taxon
3 (1954) 114.
Radermachia Thunb., Kongl. Svenska Vetensk. Acad. Handl. 37 (1776) 251 (‘Rademachia’); Houtt.,
Nat. Hist. II, Pl. 11 (1779) 446.
Polyphema Lour., Fl. Cochinch. (1790) 546.
Trees, monoecious. Leaves spirally arranged (subg. Artocarpus) or distichous (subg.

Pseudojaca); lamina pinnately veined or sometimes subtriplinerved, entire or pinnately
divided, sometimes down to the midrib; stipules free, amplexicaul (subg. Artocarpus)
or lateral (subg. Pseudojaca). Inflorescences unisexual, solitary or in pairs in the leaf
axils or on short-shoots on the older wood, pedunculate (or subsessile), bracteate or
ebracteate. Staminate inflorescences spicate, ellipsoid to cylindrical to clavate to sub-
globose-capitate; flowers numerous, free; perianth tubular, subentire or 2 – 4-lobed to
-parted; stamen 1. Pistillate inflorescences (sub)globose to ellipsoid-capitate, flowers
numerous, with the upper parts of the perianth (or entirely) fused with adjacent perianths
and/or interfloral bracts; perianth tubular with a narrow aperture; ovary free, stigma
1 or 2 of equal or unequal length. Infructescences consisting of a solid outer layer with
a smooth areolate surface or with various protuberances, a soft, more or less fleshy
middle layer containing the fruits, and a solid central part; fruit free, with a charta-
ceous to coriaceous to crustaceous pericarp or endocarp; seed without endosperm, em-
bryo straight, longitudinally to obliquely aligned, cotyledons equal or unequal, radicle
minute. — Fig. 16.
DISTRIBUTION
The genus comprises c. 45 species and ranges from Sri Lanka to South China and
through Malesia to the Solomon Islands and Australia; 32 of these species occur as
wild trees in Malesia, A. heterophyllus only as cultivated species. These species are
elements of forest in ever wet climates or where there is a short dry season, mostly
below 1000 m. Some species can be found at altitudes up to 1500 or to 1800 m, and in
New Guinea up to 2000 m.
The number of species of the genus represented in the Sino-Himalayan region is
uncertain (see p. 94 and p. 107). At most three species, including A. heterophyllus, are
endemic to the western part of the Indian Peninsula and Sri Lanka. Three species (if
not also the wild form of A. altilis) extend through New Guinea to the Solomon Islands.
The only species found in Australia is the widespread A. glaucus.
MORPHOLOGY
Habit — The trees of several species can become very tall and the trunks then often
have buttresses. The leafy twigs are slender (leptocladous) to thick (pachycladous),
e
b
a
c
f
g
i
j q
l
p m
Fig. 16. Species of Artocarpus J. & G. Forst., the surface of infructescences. a. A. anisophyllus; b.
A. lanceifolius; c. A. brevipedunculatus; d. A. odoratissima; e. A. rigidus; f, h, i. A. debilis; g. A. trecu
lianus; j. A. lowii; k. A. teijsmannii; l. A. elasticus; m. A. sericicarpus; n. A. tamaran; o, p. A. kemando;
q. A. sepicanus (after F.M. Jarrett, J. Arnold Arbor. 40 (1959) 367; all approx. × 2.3).
the latter type in combination with large leaves and infructescences that can become
large as in A. altilis. Two species with more or less large infructescences, A. hetero
phyllus and A. integer, are leptocladous but cauliflorous, with the inflorescences in
short branchlets on the trunk or main branches. Most species are evergreen and show
continuous growth. In subg. Pseudojaca some species have intermittent growth, as
described below (p. 108).
Leaves — They are spirally arranged in subg. Artocarpus and distichous in subsp.
Pseudojaca. At germination the first leaves may be opposite or they may be reduced to
scale-leaves. In the juvenile state the lamina is pinnately lobed to more deeply incised.
In some species (of subg. Artocarpus) the lamina is (also) incised in the adult state, in
A. anisophyllus usually down to the midrib and the segments petiolulate, thus the lamina
being seemingly compound.
The two subgenera do not only differ in the arrangement of the leaves but also in
the presence of glandular spots or glands on the lamina as described below for subg.
Pseudojaca (see p. 108), and in features of the stipules.
Stipules — The stipules are free and in subg. Artocarpus fully amplexicaul and rela-
tively large, longer than 1 cm, up to 25(– 35) cm in A. altilis. In subg. Pseudojaca, the
stipules are lateral and short, mostly 0.2 – 0.5 cm long, or if subpersistent on opening
shoots, then sometimes up to 1 cm long; in species with intermittent growth they may
be subpersistent, forming terminal buds.
Inflorescences — Usually solitary and axillary, but in particular staminate inflores-

cences occur on short-shoots on the older wood (ramiflorous) in some species of subg.
Pseudojaca. Cauliflory is found in subg. Artocarpus.
The flower-bearing parts vary from spikes to clavate spadices or narrowly ellipsoid
to ellipsoid or globose heads; considerable variation can be found in the same species.
The flowers are densely set and free in the staminate inflorescences and (partly) fused
in the pistillate ones. The flowers are intermixed with bracts.
Bracts — Interfloral bracts are basically peltate, but often modified, as described under
the subgenera. The transformation of bracts from the basically peltate state into various
other states is also found in the African genus Treculia (see Berg, Bull. Jard. Bot. Belg.
47 (1977) 380). Interfloral bracts, in particular peltate ones, can be caducous, in certain
species consistently so, or not. Presence or absence of interfloral bracts is rather errati-
cally distributed within the genus, its subdivisions, and even in species.
In several species, the flower-bearing part can be subtended by small basally attached
scale-like bracts, not essentially different from the inflorescences of Parartocarpus.
Pistillate flowers — The lower parts of the perianths of the pistillate flowers surround-
ing the ovaries are free. More apically the perianths are connate, forming the solid
‘skin’ of the infructescence (see Jarrett 1976). The uppermost parts of the perianths are
free, become more or less hardened and (partly) form the varied ‘ornamentation’ of the
surface of the infructescence, in particular of those of subg. Artocarpus (see Fig. 16).
The free apices of the perianths remain mostly flat in subg. Pseudojaca, but in some
species they may become low-pyramidate and hardened.
The style runs through a very narrow channel (see Fig. 16d – f). In subg. Artocarpus
the stigmas are usually either bifid (with equally or unequally long arms), or simple,
but both states can be found in the same species. In subg. Pseudojaca the stigmas are
simple with exception of those of A. altissimus.
The style remains terminal as in subg. Pseudojaca, some species of subg. Artocar
pus, those included in sect. Duricarpus by Jarrett (1959), whereas in the other species,
those included in sect. Artocarpus, the style becomes lateral to subbasal during seed-
setting.
Fruits — Ellipsoid (to globose), with a ± crustaceous to cartilaginous pericarp or endo

carp varying in size, being either (0.6 –)0.8 –1.2(–1.5) or c. 2 – 3 cm long. Large seeds
are associated with the cultivated form of the species.
Seeds — They are longitudinally aligned and with equal cotyledons in the groups with
terminal styles and oblique and with unequal cotyledons in the groups in which the
style becomes lateral to subbasal.
Seedlings — The features of seedlings of several species have been described by Burger
(1972) and Troup (1921); the germination is hypogeal; the seeds are not dormant.
Leaf anatomy — Leaf anatomical features, including the variation in pluricellular (or
glandular) trichomes, are described by Renner (1907) and Jarrett (1959); none of them
has detected the Ficus-like glandular spots in subg. Pseudojaca (see p. 108).
References: Burger, D., Seedlings of some tropical trees and shrubs mainly of South East Asia
(1972). Pudoc, Wageningen. — Jarrett, F.M., Studies in Artocarpus and allied genera, I. J. Arnold
Arbor. 40 (1959) 1– 29. — Jarrett, F.M., The syncarp of Artocarpus – a unique biological phenomenon.
Gard. Bull. Singapore 29 (1976) 35 – 39. — Renner, O., Beiträge zur Anatomie und Systematik der Ar-
tocarpeen und Conocephaleen, insbesondere der Gattung Ficus. Bot. Jahrb. Syst. 39 (1907) 319 – 448.
— Troup, R.S., The silviculture of Indian trees 3 (1921). Clarendon Press Oxford.
CHROMOSOME NUMBERS
A chromosome number of 2n = 56 has been reported for the species (Fedorov 1969;
Chen 1993; Oginuma & Tobe 1995), but the seedless (= sterile) variety of A. altilis is
hexaploid and has 2n = 84 chromosomes (see Ragone 2001).
References: Chen., R.-Y (ed.), Chromosome atlas of Chinese fruit trees and their close wild relatives.
Chromosome atlas of Chinese principal economic plants (1993). International Academic publishers,
Beijing. — Fedorov, A.A. (ed.), Chromosome numbers of flowering plants (1969). St. Petersburg.
— Oginuma, K. & H. Tobe, Karyomorphology of some Moraceae and Cecropiaceae (Urticales). J.
Pl. Res. 108 (1995) 313 – 326. — Ragone, D., Chromosome numbers and pollen stainability of three
species of Pacific island breadfruit (Artocarpus, Moraceae). Amer. J. Bot. 88 (2001) 693 – 696.
SUBDIVISION AND DELIMITATION
The genus comprises two markedly distinct subgenera: Artocarpus and Pseudojaca.
The differences are such that these two entities and Parartocarpus could be considered
as equivalent at the subgeneric rank. The latter is distinct in the basally attached bracts,
larger than those usually found in the two subgenera, and differs from subg. Artocarpus
in the small connate stipules, and from subg. Pseudojaca in the intrapetiolar position
of the connate stipules. Prainea is also very close to Artocarpus, differing in the free
pistillate flowers.
Literature: Corner, E.J.H., Notes on the systematy and distribution of Malayan phanerogams, II.
The jack and the chempedak. Gard. Bull. Singapore 10 (1938) 56 – 81. — Corner, E.J.H., Wayside
Trees of Malaya ed. 1, 1 (1940). Government Printing Office, Singapore. — Corner, E.J.H., Wayside
Trees of Malaya ed. 3, 1 (1988). Malayan Nature Society, Kuala Lumpur. — Jarrett, F.M., Studies in
Artocarpus and allied genera, III. J. Arnold Arbor. 40 (1959) 113 –155. — Van der Pijl, L., On the flower
biology of some plants from Java – with general remarks on fly-traps (species of Annona, Artocarpus,
Typhonium, Gnetum, Arisaema and Abroma). Ann. Bogor. 1 (1953) 77– 99.
DISPERSAL
Seeds are mainly dispersed by various arboreal animals such as monkeys, squirrels,
and civet cats. Monkeys and squirrels bite the infructescence into pieces and carry off
the seeds. Yellow, red, or purple infructescences are also eaten by birds. The large in-
fructescences of the cauliflorous species, A. integer and A. heterophyllus, particularly
in the latter species which can reach considerable dimensions, are probably dispersed
by forest ungulates including elephants (Ridley 1930).
References: Ridley, H.N., The dispersal of plants throughout the world (1930). Reeve & Co. Ltd.,
Ashford.
USES
The genus includes two commonly cultivated trees as fruit trees throughout the tropics:
A. altilis (Breadfruit) and A. heterophyllus (Jack or Jackfruit); A. integer (Chempedak)
is widely grown in Malesia and A. odoratissimus (Marang) is in cultivation in the Philip-
pines and Borneo (see for these species Rajendran, Soepadmo, Jansen, and Dela Cruz,
respectively, Prosea 2 (1991) 83 – 96). The infructescences of several other species,
not in cultivation, are harvested for the juicy perianth parts and /or the seeds which are
often eaten roasted (see Seibert & Jansen 1991).
The lightweight hardwood of several Artocarpus species is used for several purposes
such as for construction, veneer, and various utensils (see Djarwaningsih et al. 1995).
Several Artocarpus species also produce exudates such as latex (see Boer & Ella 2000)
to use as birdlime or as medicine or stimulants (see Van der Vossen & Wessel 2000)
and/or (bark) fibres to make cloth (see Brink & Escobin 2003).
References: Boer, E. & A.B. Ella (eds), Plant resources of South-east Asia 18. Plants producing
exudates (2000) 139 –140. Backhuys Publishers, Leiden. — Brink, M. & R.P. Escobin (eds), Plant
resources of South-east Asia 17. Fibre plants (2003) 304 – 305. Pudoc, Wageningen. — Dela Cruz,
F.S., Artocarpus odoratissimus, in: E.W.M. Verheij & R.E. Coronel (eds), Plant resources of South-east
Asia 2, Edible fruits and nuts (1991) 94 – 96. — Djarwaningsih, T., D.S. Alonzo, S. Sudo & M.S.M.
Sosef, Artocarpus J.R. & G. Forster, in: R.H.M.J. Lemmens, I. Soerianegara & W.C. Wong (eds), Plant
resources of South-east Asia 5, 2, Timber trees: Minor commercial timbers (1995) 59 –71. Pudoc, Wage-
ningen. — Jansen, P.C.M. 1991. Artocarpus integer, in: E.W.M. Verheij & R.E. Coronel (eds), Plant
resources of South-east Asia 2, Edible fruits and nuts (1991) 91– 94. Pudoc, Wageningen. — Rajendran,
R., Artocarpus altilis, in: E.W.M. Verheij & R.E. Coronel (eds), Plant resources of South-east Asia 2,
Edible fruits and nuts (1991) 83 – 86. Pudoc, Wageningen. — Seibert, B. & P.C.M. Jansen, Artocarpus
J.R. & G. Forster, in: E.W.M. Verheij & R.E. Coronel (eds), Plant resources of South-east Asia 2, Ed-
ible fruits and nuts (1991) 79 – 83. Pudoc, Wageningen. — Soepadmo, E., Artocarpus heterophyllus,
in: E.W.M. Verheij & R.E. Coronel (eds), Plant resources of South-east Asia 2, Edible fruits and nuts
(1991) 86 – 91. Pudoc, Wageningen.— Van der Vossen, H.A.M. & M. Wessels (eds), Plant resources
of South-east Asia 16. Stimulants (2000) 139. Pudoc, Wageningen.
TAXONOMIC HISTORY
For the complex taxonomic history of the genus, in particular with regard to species
in cultivation and the confusion concerning the names of these species see Jarrett, J.
Arnold Arbor. 40 (1959) 115 –125, and for an account on the nomenclatural problems
related to Artocarpus integer see Corner, Gard. Bull. Singapore 10 (1938) 56 – 81.
DUBIOUS NAMES
For dubious names and nomina nuda linked to the Malesian region see Jarrett, J. Arnold
Arbor. 41 (1960) 139 –140.
KEY TO THE SUBGENERA
1a. Leaves spirally arranged; stipules fully amplexicaul, 1 cm or longer . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subg. Artocarpus
b. Leaves distichous; stipules lateral, shorter than 1 cm . . . . . . . . Subg. Pseudojaca
Subgenus Artocarpus
Artocarpus J. & G. Forst. subg. Artocarpus: F.M. Jarrett, J. Arnold Arbor. 40 (1959) 129. — Artocarpus
J. & G. Forst. subg. Jaca Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 110. — Artocarpus J. & G.
Forst. sect. Jaca (Trécul) Renner, Bot. Jahrb. Syst. 39 (1907) 363.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Duricarpus F.M. Jarrett, J. Arnold Arbor. 40 (1959)
137.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Duricarpus F.M. Jarrett ser. Laevifolii F.M. Jarrett,
J. Arnold Arbor. 40 (1959) 138.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Duricarpus F.M. Jarrett ser. Asperifolii F.M. Jarrett,
J. Arnold Arbor. 40 (1959) 143.
Artocarpus J. & G. Fost. subg. Artocarpus sect. Artocarpus ser. Incisifolii F.M. Jarrett, J. Arnold
Arbor. 40 (1959) 298.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Artocarpus ser. Cauliflori F.M. Jarrett, J. Arnold
Arbor. 40 (1959) 327.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Artocarpus ser. Angusticarpi F.M. Jarrett, J. Arnold
Arbor. 40 (1959) 338.
Artocarpus J. & G. Forst. subg. Artocarpus sect. Artocarpus ser. Rugosi F.M. Jarrett, J. Arnold Arbor.
40 (1959) 343.
Leaves spirally arranged; lamina without glands or glandular spots; stipules fully
amplexicaul, usually longer than 1 cm. Staminate flowers with largely connate te-
pals. Pistillate flowers with cushion shaped, pyramidate, conical, spine-like, subulate,
or filiform free apices, indurated in fruit. Flowers often intermixed with peltate to
spathulate (in pistillate inflorescences often (partly) caducous) bracts, or in staminate
inflorescences with rod-shaped interfloral processes, and in pistillate inflorescences
with conical, subulate or filiform processes, or else bracts and processes absent.
DISTRIBUTION
This subgenus is concentrated in the western part of the Malesian region with 19 in-
digenous species, some species extending to the Asian mainland and some species to
western New Guinea. Artocarpus altilis is clearly eastern Malesian, extending into
Micronesia, Melanesia, and Polynesia, mainly with cultivars. Artocarpus sepicanus is
endemic to New Guinea and A. teijsmannii extends with its subsp. subglabrus to the
Solomon Islands.
Artocarpus nobilis Thwaites is endemic to Sri Lanka. The origin of A. heterophyl
lus lies probably in the western Indian Peninsula. Clear morphological connections of
A. hirsutus with material from the Sino-Himalayan region (as indicated below under
A. hispidus, see p. 93) makes it uncertain whether A. hirsutus is a distinct species and
another endemic of the Indian Peninsula.
MORPHOLOGY
Interfloral bracts — In staminate inflorescences, the bracts are basically peltate (to
spathulate), but may be modified into rod-shaped ‘processes’, which were described
by Jarrett as sterile flowers.
In pistillate inflorescences lacking (sub)peltate bracts, the flowers are often inter-
mixed with rigid ‘processes’ varying in shape and length from conical to filiform, and
from straight to ± curved. These are apparently also modified bracts. These ‘processes’
often resemble the hardened free apices of the perianths which vary from cushion
shaped to pyramidate to conical to spine-like to subulate to filiform (and straight to
curved). The perianth apices can be distinguished from ‘processes’ by the perforate
apex or also by the persisting stigmata. The variation of the apices of perianths and the
interfloral ‘processes’ is illustrated in Fig. 16.
SUBDIVISION
A group of species included by Jarrett (1959) in sect. Duricarpus is characterised

by globose (to ellipsoid) infructescences with indurated apices ranging from cushion
shaped to (up to 1 cm long) cylindrical to spine-like to clavate (persistent or cadu-
cous) peltate interfloral bracts, terminal styles and longitudinally aligned embryos with
equal cotyledons. This group comprises A. anisophyllus, A. annulatus, A. asperula
Gagnep. (Indochina), A. brevipedunculatus, A. calophylla Kurz (Myanmar), A. chama
Buch.-Ham. (Sikkim to Vietnam), A. hirsutus Lam. (India), A. hispidus, A. lanceifolius,
A. nobilis Thwaites (Sri Lanka), A. odoratisssimus, A. rigidus, and A. sarawakensis
(but see p. 93). Artocarpus hirsutus deviates in the pendulous spicate staminate inflo-
rescences and in the absence of peltate interfloral bracts.
A second main group (section) comprises species included in sect. Artocarpus which
is characterised by the frequent absence of peltate interfloral bracts, being substituted
by elongate processes, or totally absent. The apices of the perianths varies from cushion
shaped to spine-like. The style is lateral to subbasal. The embryos are obliquely aligned
with unequal to equal cotyledons. The group can be subdivided into four subgroups:
1) Plants with cauliflorous species: A. heterophyllus and A. integer.
2) Plants usually with incised laminas in the adult state: A. altilis and possibly also
A. treculianus in which the lamina is sometimes pinnately incised in the adult state.
The latter species also shows affinities to A. teijsmannii.
3) Plants with entire laminas in the adult state, the staminate inflorescences with
smooth surfaces and the vegetative parts inconspicuously whitish (to brownish)
hairy or glabrous: A. excelsus, A. lowii, A. sepicanus, and A. teijsmannii.
4) Plants with entire laminas in the adult state, the staminate inflorescences with
sulcate to tuberculate surfaces and the vegetative parts more or less conspicuously
brown hairy: A. elasticus, A. kemando, A. sericicarpus, and A. tamaran.
KEY TO THE SPECIES OF SUBGENUS ARTOCARPUS
1a. Inflorescences borne on stems and/or branches (cauli- or ramiflorous) . . . . . . . . 2

b. Inflorescences axillary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Indumentum whitish; peduncle broadened into a rim. — Widespread, in cultiva-
tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. A. heterophyllus
b. Indumentum brownish or absent; peduncle not broadened into a rim. — Wide-
spread, indigenous or in cultivation . . . . . . . . . . . . . . . . . . . . . . . . . 9. A. integer
3a. Lamina deeply pinnately incised with 12 – 20 segments on each side, the incisions
down to near the midrib. — Philippines . . . . . . . . . . . . . . . . . . . . . . . 1. A. altilis
b. Lamina entire or pinnately incised, if deeply so and with up to 12 segments on each,
then these mostly petiolulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Staminate and pistillate inflorescences with peltate interfloral bracts . . . . . . . . . . 5
b. Staminate and pistillate inflorescences without interfloral bracts, or if present, then
not peltate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
5a. Epidermis of petiole flaking off. — New Guinea . . . . . . . . . . . 17. A. sepicanus
b. Epidermis of petiole persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Lateral veins 4 – 6 pairs; uncinate hairs on leafy twigs, petioles, and margin of the
lamina (or glabrous). — Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . 3. A. annulatus
b. Lateral veins 6 – 20 pairs; straight and persistent hairs on leafy twigs, petioles, and
margin of the lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Lamina of adult specimens incised down to the midrib, the segments often petiolu-
late. — Malay Peninsula, Sumatra, Borneo (and Philippines?) . 2. A. anisophyllus
b. Lamina of adult specimens entire or, if pinnately incised, then not down to the
midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Lamina of adult specimens usually pinnately incised; petiole 3 – 9(–13); stipules
usually 10 –15 cm long. — Widespread . . . . . . . . . . . . . . . . . . . . . . . . 1. A. altilis
b. Lamina of adult specimens entire (or 3-lobed to -fid); petiole 1– 3(– 4) cm long;
stipules 1– 5(– 8) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9a. Leafy twigs and stipules glabrous. — New Guinea . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20b. A. teijsmannii subsp. subglabrus
b. Leafy twigs and stipules hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Hairs on the stipules longer at the apex (subsericeous) than lower down (appress-
edly puberulous). — Widespread . . . . . . 20a. A. teijsmannii subsp. teijsmannii
b. Hairs on the stipules equally long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11a. Lamina glabrous and both surfaces smooth. — Malay Peninsula, Sumatra, Bor-
neo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. A. lanceifolius
b. Lamina hairy, at least beneath on the main veins, often scabrous or scabridulous
above and/or beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Hairs on the leafy twigs appressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
b. At least some hairs on the leafy twigs ± patent . . . . . . . . . . . . . . . . . . . . . . . . . 15
13a. Stipules 0.5 –1 cm long; lateral veins usually 7–10 pairs; lamina smooth above.
— Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. A. brevipedunculatus
b. Stipules 1– 5(– 8) cm long; lateral veins usually 10 – 20 pairs; lamina ± scabrous
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a. Peduncle of the pistillate inflorescences 5 –14 cm long and the free parts of the
perianth clavate, 8 –12 mm long; head of staminate inflorescences 4 – 9 cm long.
— Borneo, Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. A. odoratissimus
b. Peduncle of the pistillate inflorescences 0.8 – 4 cm long and the free parts of the
perianth cylindrical, 3 – 8 mm long; head of staminate inflorescences 1.5 – 3 cm
long. — Sumatra, Malay Peninsula, Java, Borneo, Lesser Sunda Islands, Malay
Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. A. rigidus
15a. Lamina smooth above. — Sumatra, Borneo . . . . . . . . . . . . 16. A. sarawakensis
b. Lamina ± scabrous above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16a. Stipules 0.5 –1(– 2) cm long and petioles 1–1.5(– 2) cm long; peduncle of the
staminate inflorescences 1– 3 cm long; peduncle of the pistillate inflorescences
2.5 – 3.5 cm long. — Malay Peninsula . . . . . . . . . . . . . . . . . . . . . 8. A. hispidus
b. Stipules 1– 5(– 8) cm long and petioles (1.5 –)2 – 3 cm long; peduncle of the stami-
nate inflorescences usually 3 –13 cm long; peduncle of the pistillate inflorescences
5 –14 cm long. — Borneo, Philippines . . . . . . . . . . . . . . . . 14. A. odoratissimus
17a. Surface of the staminate inflorescences sulcate or tuberculate, or the flowers inter-
mixed with subulate processes and/or the pistillate inflorescences with numerous
subulate to filiform interfloral processes, these at least 2 times longer than the free
parts of the perianths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
b. Staminate inflorescences with a smooth surface and the pistillate inflorescences
without interfloral processes or if processes present, then few and less than 2 times
as long as the perianths, or else c. 3 times as long as the perianths and pyrami-
date . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
18a. Lateral veins (13 –)18 – 24 pairs, rarely branched or forked away from the margin;
stipules 3 – 9 cm long; petiole 3.5 – 4 cm long. — Borneo . . . . . . 19. A. tamaran
b. Lateral veins (6 –)10 –16(–18) pairs, most or some of them branched or forked
away from the margin, or if rarely branching, then the stipules 0.7– 2 cm long and
petiole 1– 3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19a. Leafy twigs 2 – 3 mm thick; stipules 0.7– 2 cm long; lateral veins rarely branched
or forked away from the margin; peduncle of staminate inflorescences 0.3 – 2 cm
long; peduncle of pistillate inflorescences 0.3 – 0.8 or 1– 4 cm long, brown veluti-
nous. — Malay Peninsula, Sumatra, Borneo . . . . . . . . . . . . . . . . 10. A. kemando
b. Leafy twigs 2 – 20 mm thick; stipules mostly longer than 2 cm; lateral veins mostly
branched or forked away from the margin; peduncle of staminate inflorescences
usually more than 2 cm long, and of pistillate inflorescences more than 4 cm long
or if less than 4 cm long, then minutely whitish puberulous . . . . . . . . . . . . . . . 20
20a. Hairs on vegetative parts minute to short, appressed and white, except for the
brown subsericeous apex of the stipules; staminate inflorescences with subulate
interfloral processes. — Widespread . . . 20a. A. teijsmannii subsp. teijsmannii
b. Hairs on vegetative parts predominantly brown, on the stipules not distinctly dif-
ferent in length and colour; staminate inflorescences without subulate interfloral
processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Hairs on petioles and peduncles ± patent; indumentum on processes of the pistil-
late inflorescences partly hirtellous. — Borneo, Philippines, Celebes, Moluccas
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. A. sericicarpus
b. Hairs on petioles and peduncles appressed; indumentum on processes of the pistil-
late inflorescences puberulous to hispidulous. — Sumatra, Malay Peninsula, Java,
Borneo, Philippines (Palawan), Lesser Sunda Islands . . . . . . . . . 5. A. elasticus
22a. Petiole 3 –7(–10) mm thick, (2.5 –)3.5 –10 cm long; stipules often longer than 10
cm and brown hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
b. Petiole 1– 3 mm thick, mostly 1– 3.5 cm long, or if up to 5 cm long, then the hairs
on the stipules appressed; stipules up to 8 cm long . . . . . . . . . . . . . . . . . . . . . . 24
23a. Lamina of adult specimens mostly pinnately incised; areoles of lamina flat be-
neath. — Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. A. altilis
b. Lamina of adult specimens usually entire; areoles of lamina bullate beneath.
— Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. A. elasticus
24a. Lateral veins 7–10 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
b. Lateral veins 10 – 20 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
25a. Hairs on the stipules becoming longer towards the apex. — Widespread . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20a. A. teijsmannii subsp. teijsmannii
b. Hairs on the stipules about equally long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26a. Indumentum on leafy twigs predominantly brown(ish); petiole 1– 2 mm thick.
— Sumatra, Malay Peninsula, Borneo . . . . . . . . . . . . . . . . . . . . 10. A. kemando
b. Indumentum on leafy twigs predominantly whitish; petiole 2 – 3 mm thick. — Bor-
neo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. A. obtusus
27a. Petioles and peduncles with (longer) uncinate hairs. — Malay Peninsula . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. A. lowii
b. Petioles and peduncles without uncinate hairs . . . . . . . . . . . . . . . . . . . . . . . . . 28
28a. Stipules 6 –18 cm long; petiole 3 – 8 cm long. — Philippines . 21. A. treculianus
b. Stipules usually 0.7– 5 cm long; petiole 1– 3.5 cm long . . . . . . . . . . . . . . . . . . 29
29a. Stipules 0.7– 2 cm long; petiole 1– 2 mm thick; indumentum predominantly
brown(ish). — Sumatra, Malay Peninsula, Borneo . . . . . . . . . . 10. A. kemando
b. Stipules usually 2 – 6 cm long; petiole usually 2 – 3 mm thick, or if 1– 2 mm thick,

then the indumentum predominantly whitish . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Hairs of the stipules becoming longer towards the apex. — Widespread . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20a. A. teijsmannii subsp. teijsmannii
b. Hairs of the stipules about equally long — Borneo. . . . . . . . . . . . . 6. A. excelsus
1. Artocarpus altilis (Parkinson) Fosberg

Artocarpus altilis (Parkinson) Fosberg, J. Wash. Acad. Sci. 31 (1941) 95; Brittonia 12 (1960) 104;
Backer & Bakh.f., Fl. Java 2 (1965) 18; R. Rajendran, Prosea 2 (1991) 83, cum t. — Sitodium altile
Banks & Sol. ex Parkinson, J. Voy. South Seas (1773) 45.
Artocarpus communis J. & G. Forst., Char. Gen. Pl., ed. 2 (1776) 101, t. 51, 51a; King, Ann. Roy. Bot.
Gard. (Calcutta) 2 (1889) 16; Saff., Contr. U.S. Natl. Herb. 9 (1905) 189, t. 7, 27, 36; Merr., Philipp.
J. Sci., 1, Suppl. (1906) 43; Fl. Manila (1912) 176; Interpr. Herb. Amboin. (1917) 190; Sp. Blancoan.
(1918) 123; W.H. Br., Min. Prod. Philip. For. 2 (1921) 162, t. 13, 14; Merr., Enum. Philipp. Flow.
Pl. 2 (1923) 40; Ochse & Bakh., Veg. Dutch East Indies (1931) 488; H.J. Lam, Blumea 1 (1934)
119; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 250; Brass & A.L. Rand, Bull. Amer. Mus.
Nat. Hist. 77 (1940) 366, 367; Quisumb., Philipp. J. Sci. 72 (1940) 331; Archbold, A.L. Rand &
Brass, Bull. Amer. Mus. Nat. Hist. 79 (1942) 233, 235; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 307;
Kochummen, Tree Fl. Malaya 3 (1978) 124.
Radermachia incisa Thunb., Kongl. Svenska Vetensk. Acad. Handl. 37 (1776) 253; Houtt., Nat. Hist.
II, Pl. 11 (1779) 449, t. 76. — Sitodium incisum (Thunb.) Thunb., Philos. Trans. 69 (1779) 465.
— Artocarpus incisus (Thunb.) L.f., Suppl. (1781) 411; G. Forst., Vom Brodbaum (1784); Blume,
Bijdr. (1825) 480; Hook., Bot. Mag. 55 (1828) t. 2869 – 2871; Roxb., Fl. Ind., ed. Carey 3 (1832)
527; Decne., Herb. Timorensis Descr. (1835) 164; Blanco, Fl. Filip. (1837) 668; Span., Linnaea 15
(1841) 343; Blanco, Fl. Filip., ed. 2 (1845) 465; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 110;
Miq., Pl. Jungh. (1851) 44; Fl. Ned. Ind. 1, 2 (1859) 285; Fl. Ned. Ind., Eerste Bijv. (1861) 417;
Baill., Adansonia 4 (1863) 79, t. 5; Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211; Blanco,
Fl. Filip., ed. 3, 3 (1879) 75; Fern.-Vill., Nov. App. (1880) 202; S. Vidal, Revis. Pl. Vasc. Filip.
(1886) 254; K. Schum., Bot. Jahrb. Syst. 9 (1888) 199; Warb., Bot. Jahrb. Syst. 13 (1891) 295;
K. Schum., Notizbl. Bot. Gart. Berlin-Dahlem 2 (1898) 110; K. Schum. & Lauterb., Fl. Schutzgeb.
Südsee (1901) 267; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 15; Koord., Exkurs.-Fl.
Java 2 (1912) 92; Ridl., Fl. Malay Penins. 3 (1924) 351; Lane-Poole, For. Res. (1925) 81; Lauterb.,
Bot. Jahrb. Syst. 63 (1930) 421, 437, 442; Corner, Gard. Bull. Singapore 10 (1939) 290; Wayside
Trees Malaya (1940) 655, t. 196; H. St. John, Pacific Sci. 2 (1948) 109; Corner, Wayside Trees
Malaya ed. 3, 2 (1988) 518, t. 161. — Artocarpus incisifolia Stokes, Bot. Mat. Med. 4 (1812) 331,
substitute name.
Artocarpus camansi Blanco, Fl. Filip. (1837) 666; ed. 2 (1845) 467; ed. 3, 3 (1880) 76; Merr., Sp.
Blancoan. (1918) 124; Quisumb., Philipp. J. Sci. 72 (1941) 331; Quisumb., Med. Pl. Philipp.
(1951) 226.
Artocarpus rima Blanco, Fl. Filip. (1837) 671; ed. 2 (1845) 467; ed. 3, 3 (1880) 77; Merr., Sp. Blan-
coan. (1918) 123.
Artocarpus laevis Hassk., Flora 25, 2 (1842) Beibl. 18; Trécul, Ann. Sci. Nat., Bot. sér 3, 8 (1847)
122. — Artocarpus incisus (Thunb.) L.f. var. laevis (Hassk.) Miq., Pl. Jungh. (1851) 44; Fl. Ned.
Ind. 1, 2 (1859) 285.
Artocarpus mariannensis Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 114; Fosberg, Brittonia 12 (1960)
108.
Artocarpus incisus (Thunb.) L.f. var. muricata Becc., For. Borneo (1902) 108.
Artocarpus pinnatisectus Merr., Philipp. J. Sci. 18 (1921) 50; Enum. Philipp. Flow. Pl. 2 (1923) 43;
F.M. Jarrett, J. Arnold Arbor. 40 (1959) 323, t. 13.
Artocarpus blancoi (Elmer) Merr., Enum. Philipp. Flow. Pl. 2 (1923) 40; Quisumb., Med. Pl. Philipp.
(1951) 226; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 301. — Artocarpus communis J. & G. Forst.
var. blancoi Elmer, Leafl. Philipp. Bot. 2 (1909) 617.
Artocarpus communis J. & G. Forst. var. pungens J.J. Sm. ex K. Heyne, Nutt. Pl. Ned.-Ind., ed. 2
(1927) 557.
Artocarpus leeuwenii Diels, Bot. Jahrb. Syst. 67 (1935) 173.
Artocarpus horridus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 306.
Artocarpus multifidus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 324, t. 13.
Artocarpus elasticus auct. non Blume: Hassk., Abh. Naturf. Ges. Halle 9 (1866) 158; Merr., Interpr.
Herb. Amboin. (1917) 191.
Artocarpus papuana auct. non Renner: Diels, Bot. Jahrb. Syst. 67 (1935) 175.
[Soccus lanosus Rumph., Herb. Amboin. 1 (1741) 110, t. 32.]
[Soccus granosus Rumph., Herb. Amboin. 1 (1741) 112, t. 33.]
[Soccus sylvestris Rumph., Herb. Amboin. 1 (1741) 114, t. 34.]
Tree up to 35(– 40) m tall, with buttresses, evergreen or deciduous. Leafy twigs 5 – 22
mm thick, minutely whitish appressedly puberulous to subglabrous, often also brown
(to yellowish) (sub)hirsute or setose (to strigose), drying brown to blackish; lenti-
cels concentrated in the upper part of the internodes or ± scattered, few or numerous.
Leaves spirally arranged; lamina coriaceous to chartaceous, elliptic to subrhombic to
obovate in outline, 20 – 80(–100) by 15 – 50(– 60) cm, mostly pinnatifid with 1– 5(– 9,
occasionally – 20) pairs of lateral lobes or segments, the incisions mostly to halfway,
occasionally down to near the midrib, base cuneate to rounded (to subcordate), apex
short-acuminate to acute, margin subentire to repand (to coarsely dentate), ± revolute;
upper surface whitish to brownish (appressedly) puberulous to hirtellous or to sub-
hispidulous, also brown set(ul)ose to strig(ill)ose on the main veins (or subglabrous),
smooth to scabridulous; lower surface whitish to brownish appressedly puberulous (or
to subhirtellous) on the veins, the hairs straight, on the smaller veins the hairs patent
and all or some uncinate, all hairs on the smaller veins minute and appressed, or on the
main veins also brown set(ul)ose, strig(ill)ose, or subhirsute, scabridulous to smooth;
lateral veins 9 –13(–18) pairs, (in entire laminas) some or most branched or forked away
from the margin, tertiary venation scalariform, ± prominent; petiole 3 – 9(–13) cm long,
3 –7(–10) mm thick, whitish to brownish appressedly puberulous, often also brown
(sub)hirsute or set(ul)ose (or strigillose); stipules (3 –)10 – 25(– 35) cm long, whitish to
brownish appressedly puberulous, often also brownish (sub)strigillose to (sub)hirsute
or setose (to strigose), or brownish to yellowish subsericeous to subvillous or hirsute,
caducous. Staminate inflorescences axillary, solitary; peduncle 1– 6 cm long, whitish
to brownish appressedly puberulous to substrigillose or to hirtellous, or also brown
(sub)hirsute or setulose (to strigose), ± scabrous to smooth; head cylindrical to clavate
to spicate, 5 – 30(– 40) by (0.5 –)1– 3(– 5.5) cm; perianth tubular, 1– 2 mm long, the apex
2-lobed, subhispidulous or subglabrous; stamen 1.5 – 2.8 mm long, anther 0.6 – 0.8 mm
long; interfloral bracts absent (or present and peltate). Pistillate inflorescences axil-
lary, solitary; peduncle 2 –14 cm long, whitish strigillose and also brown setulose or
brown to yellowish hirtellous to (sub)hirsute, ± scabrous to smooth; head obovoid, el-
lipsoid, subglobose, or cylindrical; perianth tubular, c. 1 mm long, whitish to brownish
(retrorsely) hispidulous to puberulous (or subglabrous), the apex 2-lobed to convex;
stigma bifid or simple; interfloral bracts usually absent, sometimes present, and then
often as rod-shaped processes. Infructescences ellipsoid to subglobose to obovoid or to

cylindrical, 5 –10(–15) by 5 –10 cm or 15 – 30 by 10 – 30 cm diam., covered by 2 – 3 mm
long pyramidate, 2 – 8 mm long cylindrical, or 8 –15 mm long subulate to filiform, often
± curved apices of the perianths, sometimes intermixed with 0.5 – 2 cm long subulate
to filiform, often ± curved processes, rarely with peltate bracts; fruits ellipsoid, 1–1.2
or 2 – 2.5 cm long. — Fig. 17.
7 cm
3 cm
2 mm
0.5 mm
a 1.5 cm e c d b
Fig. 17. Artocarpus altilis (Parkinson) Fosberg. a. Leafy twig with pistillate inflorescence; b. staminate
inflorescence; c. length section of young staminate flower; d. mature staminate flower; e. cross section
of infructescence (all: Beguin 1976).
Distribution — Melanesia, Micronesia, Polynesia; in Malesia: Philippines, Celebes,

Lesser Sunda Islands (Flores), New Guinea; in these islands also the cultivated seeded
and seedless forms of the species; these are also in cultivation elsewhere in Malesia (and
throughout the tropics) (see also Barrau in Simmonds, Evolution of crop plants (1979)
202); there are some indications that the wild form extends to the Solomon Islands.
Seeded cultivars may naturalize.
Habitat — Evergreen forest at low altitudes.
Notes — 1. In Malesia, three forms can be distinguished: the wild form with rela-
tively small infructescences (up to 10 or 15 cm long) with relatively small seeds (1–1.2
cm long) and two cultivated forms with larger infructescences (up to 30 cm long or
wide), the seeded form with larger fruits (2 – 2.5 cm long). The wild form is, with regard
to the indumentum, rather variable: varying from sparse to dense on various parts (such
as the stipules), from short to long, from rather weak to rigid and pungent (or setose),
from straight to uncinate (when juvenile?), from whitish to yellowish to (dark) brown.
Material with stiff and pungent hairs was included in A. horridus (by Jarrett 1959) and
is the common form in the Moluccas and western New Guinea. The wild form also
shows a considerable variation in shape and length of the apices of the perianths of the
pistillate flowers, varying in length from 2 to 15 mm, and in shape from pyramidate to
filiform. Plants with the longer perianths (10 –15 mm) and filiform apices are mainly
found in the Philippines (but do occur elsewhere, e.g., in Flores). These have been in-
cluded in A. blancoi (by Jarrett 1959), but perianth with short-tubular apices also occur
in Luzon and Palawan. The stigmas are simple or bifid, short or rather long, and straight
or twisted. Lack of adequate material makes it difficult to figure out to what extent the
variation mentioned above is also found in the cultivated form(s) in Malesia and other
tropical regions where the species is in cultivation.
2. The infructescences and seeds of this species, known as the ‘breadfruit’, are (and
have been) important food sources. A detailed account on the economic importance and
history, and morphological variation of the cultivated forms was presented by Jarrett
(J. Arnold Arbor. 40 (1959) 309 – 323), Barrau (1979: 201– 202), Zerega (2003/2004),
and Zerega et al. (2004).
3. The names Artocarpus camansi and A. rima are based on vernacular names applied
in the Philippines to the seeded and seedless cultivars of the species.
4. The normal chromosome number of this species is 2n = 56, and is linked with
normal seed and pollen production. The seedless cultivars have 2n = 84 and pollen
production is poor. The situation is less clear in Micronesia (see below).
5. Molecular studies by Zerega et al. (2004) confirmed the complexity of origin and
distribution of cultivated forms of the species. The study recognizes three main groups
(treated as species): ‘A. camasi’ probably representing the material indicated as the
wild form above, ‘A. mariannensis’, a somewhat different Micronesian wild form of
the species (and its derivates in cultivation), and ‘A. altilis’, representing the diverse
cultivated material from Melanesia and Polynesia.
6. Fosberg (1960) referred the Micronesian material of this species partly to Artocar
pus mariannensis, partly to A. altilis, and the rest to hybrids between the two ‘species’.
A study by Ragone (2001) on chromosome numbers and fertility in Artocarpus material
of the Pacific also revealed partial and full sterility in tetraploid material.
7. It is with some hesitation that A. multifida and A. pinnatisectus are included, not
because of the numerous lobes of the lamina, or the anomalous staminate inflorescences
(see Jarret, J. Arnold Arbor. 40 (1959) 324), but because of the presence of the few
peltate interfloral bracts in the pistillate inflorescences. Presence or absence of inter-
floral bracts is not consistent in the subgenus and interfloral bracts (or processes) tend
to occur in material of the species from the Philippines, but not (or rarely) elsewhere
within the species range.
References: Barrau, J., Breadfruit and relatives, in: I. Simmonds (ed.), Evolution of crop plants
(1979). Longman, New York. — Fosberg, F.R., Introgression in Artocarpus (Moraceae) in Micronesia.
Brittonia 12 (1960) 101–113. — Ragone, D., Chromosome numbers and pollen stainability of three
species of Pacific breadfruit (Artocarpus, Moraceae). Amer. J. Bot. 88 (2001) 693 – 696. — Zerega,
N.J.C., The breadfruit trail, the wild ancestors of staple food illuminate human migrations. Natural His-
tory (2003/2004) 46 – 51. — Zerega, N.J.C., D. Ragone & T.J. Motley, Complex origins of breadfruit
(Artocarpus altilis, Moraceae): implications for human migrations in Oceania. Amer. J. Bot. 91 (2004)
760 –766.
2. Artocarpus anisophyllus Miq.

Artocarpus anisophyllus Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 422; Renner, Bot. Jahrb. Syst. 39
(1907) 366; Merr., Pl. Elmer. Born. (1929) 45; Corner, Wayside Trees Malaya (1940) 652, t. 191,
192; F.G. Browne, For. Trees Sar. & Brunei (1955) 352; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
138; Kochummen, Tree Fl. Malaya 3 (1978) 123, t. 2; Corner, Wayside Trees Malaya ed. 3, 2 (1988)
515, t. 155, 156; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 191, t. 2.
Artocarpus klidang Boerl., Handl. Fl. Ned. Ind. 3 (1900) 333, 371, in clavi.
Artocarpus superba Becc., For. Borneo (1902) 625; Merr., Philipp. J. Sci. 18 (1921) 51; Enum. Philipp.
Flow. Pl. 2 (1923); Ridl., Fl. Malay Penins. 3 (1924) 356.
Artocarpus anisophyllus Miq. var. sessilifolius Kochummen, Gard. Bull. Singapore 50 (1998) 200.
Tree up to 45 m tall, evergreen. Leafy twigs 10 – 20(– 40) mm thick, ± densely brown
(to whitish) appressedly puberulous to hispidulous, scabridulous or smooth, drying
brown; lenticels scattered. Leaves spirally arranged; lamina coriaceous, 30 –100(–150)
cm long, pinnately incised down to the midrib, with 5 –12 segments on each side, these
alternate or subopposite, mostly 1– 6(– 8) cm long petiolulate, subovate to ovate to
elliptic, (3.5 –)10 – 20(– 40) by (2 –)4 – 8(–13) cm, apex acuminate, base subcordate to
subcuneate, usually ± decurrent, margin subentire (or pinnately lobed to parted); upper
surface glabrous; lower surface sparsely brownish strigillose on the midrib; lateral veins
7– 20 pairs, ‘tertiary’ venation scalariform slightly prominent; ‘rachis’ sparsely brown-
ish strigillose, terminal segment often 3-lobate; petiole 4 – 9(–16) cm long, (2.5 –)5 –12
mm thick, sparsely brownish strigose, the lower part ± thickened, the epidermis per-
sistent; stipules fully amplexicaul, (2 –)4 –10(–17) cm long, brown substrigose to sub-
hirsute, caducous. Staminate inflorescences axillary, usually solitary; peduncle 3 – 6.5
cm long, brownish (to whitish) appressedly puberulous; head ellipsoid to narrowly
ellipsoid to cylindrical, 3 –7.5 by 1.5 – 2 cm; perianth tubular c. 1.8 mm long, the apex
2-lobed, minutely puberulous to subglabrous; stamen 2 – 2.5 mm long, anther c. 0.2 mm
long; interfloral bracts peltate, the apical part 0.6 – 0.8 mm diam., radially appressedly
puberulous. Pistillate inflorescences axillary, usually solitary; peduncle 5 –11 cm long,
brownish (to whitish) appressedly puberulous; head ellipsoid to subglobose; perianth
tubular, the apex convex to flat, minutely brown puberulous to muriculate, aperture
surrounded by a rim; stigma simple; interfloral bracts peltate, caducous, the apical
part 0.6 – 0.8 mm diam., radially appressedly puberulous. Infructescences subglobose,
7– 8(–10) cm diam., covered with 6 – 8 mm long, cylindrical apices of the perianths;
fruits ellipsoid, 1–1.5 cm long.
Distribution — Sumatra, Malay Peninsula, Borneo, Philippines (see note 2).
Habitat — Evergreen forest, at altitudes up to c. 1300 m.
Notes — 1. The leaf is not truly compound as there is no articulation between ‘leaflet’
and ‘rachis’. When juvenile, the lamina may not (yet) have petiolulate segments, but seg-
ments broadly attached to the ‘rachis’, a feature sometimes retained in the adult state.
2. The species is also recorded for the Philippines (Basilan, Mindanao, Palawan) by
Merrill (1921) under Artocarpus superba, but the collections were not traced by Jarrett
(1959), nor for the present treatment. Possibly specimens of A. altilis with lamina’s with
numerous incisions were misidentified as A. anisiophyllus.
3. Artocarpus annulatus F.M. Jarrett

Artocarpus annulatus F.M. Jarrett, Blumea 22 (1975) 409; Kochummen, Tree Fl. Sabah & Sarawak
3 (2000) 192.
Tree up to 25 m tall, evergreen. Leafy twigs 3 – 5 mm thick, sparsely brown puberulous

with (retrorse) uncinate hairs, glabrescent, smooth, drying brown; lenticels scattered,
few. Leaves spirally arranged; lamina subcoriaceous, entire, elliptic to obovate, 10 – 25
by 5 –16 cm, apex short-acuminate, base cuneate to obtuse, often mostly almost equal,
margin entire, ± revolute; both upper and lower surface (sub)glabrous, but initially with
brown uncinate hairs on the margin, sometimes the midrib minutely brown puberulous
beneath, smooth; lateral veins 4 – 6 pairs, some or none forked away from the margin,
tertiary venation scalariform, flat; petiole 2.5 – 5 cm long, 2 – 3 mm thick, puberulous
with brown (retrorse) uncinate hairs, glabrescent, the epidermis persistent; stipules fully
amplexicaul, 1– 2 cm long, sparsely brown hirtellous or partly puberulous with retrorse
uncinate hairs, caducous. Staminate inflorescences axillary, solitary; peduncle 1.5 – 2.5
cm long, puberulous with brown (retrorse) uncinate hairs, glabrescent; head ellipsoid
to ovoid, up to 4 by 2 cm, with annular grooves; perianth tubular, c. 0.8 mm long, the
apex 2(– 4)-lobed, minutely puberulous; stamen c. 1 mm long, anther c. 0.1 mm long;
interfloral bracts (sub)peltate, the apices 0.2 – 0.4 mm diam., puberulous. Pistillate in
florescences axillary, solitary; peduncle c. 3 cm long, puberulous with brown (retrorse)
uncinate hairs; head ellipsoid; perianth tubular, brown puberulous to subhispidulous, the
apex convex; stigma simple; flowers intermixed with interfloral bracts, these spathulate
to peltate, the apices 2 – 4 mm diam., brown puberulous. Infructescences ellipsoid, up
to 8 by 6 cm (immature), covered with 6 – 8 mm long, subulate apices of the perianths;
fruits not seen.
Distribution — Borneo (Sarawak).
Habitat — Evergreen forest at low altitudes, on limestone.
Notes — 1. The species is characterised by the small number of lateral veins and
the presence of uncinate hairs on various parts. Straight hairs are only present on the
stipules and sometimes on the midrib of the lamina beneath.
2. The infructescence is (very) fragrant.
4. Artocarpus brevipedunculatus (F.M. Jarrett) C.C. Berg

Artocarpus brevipedunculatus (F.M. Jarrett) C.C. Berg, Blumea 50 (2005) 541. — Artocarpus melinox
ylus Gagnep. subsp. brevipedunculatus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 144; Kochummen,
Tree Fl. Sabah & Sarawak 3 (2000) 202.
Tree up to 25 m tall, evergreen. Leafy twigs 3 –7 mm thick, ± densely brown strigil-

lose to hispidulous (or also hirtellous to subhirsute), often scabridulous, drying brown;
lenticels scattered. Leaves spirally arranged; lamina coriaceous, entire, elliptic to sub-
obovate, (4 –)10 – 20(– 30) by (2 –)5 – 9(–18) cm, apex short-acuminate, base cuneate
to obtuse, margin entire; upper surface sparsely brownish strigillose to subhirtellous
on the midrib (and lateral veins), smooth; lower surface brownish strigose to strigil-
lose to puberulous to hispidulous, or on the veins, ± scabrous; lateral veins (6 –)7–10
pairs, none branched or forked away from the margin, tertiary venation subscalariform
to reticulate, slightly prominent; petiole 1– 2 cm long, 2 – 2.5 mm thick, brown strig-
illose (or also hirtellous to subhirsute), the epidermis persistent; stipules 0.5 –1 cm,
brown strigillose or sometimes hirtellous, caducous. Staminate inflorescences axillary,
solitary; peduncle 3 – 3.5 cm long, brown (to whitish) appressedly puberulous; head
subglobose to ellipsoid, 1.5 – 2 by 1–1.5 cm; perianth tubular c. 1 mm long, the apex
(deeply) 2- (or 3-)lobed, minutely puberulous; stamen c. 1.5 mm long, anther c. 0.2 mm
long; interfloral bracts peltate, the apical part 0.3 – 0.4 mm diam., radially appressedly
puberulous. Pistillate inflorescences axillary, solitary; peduncle 1– 4 cm long, densely
brown appressedly puberulous; head subglobose to ellipsoid; perianth tubular, brown
puberulous, the apex ± convex, the aperture surrounded by a rim; stigma simple; inter-
floral bracts peltate, the apical part 0.3 – 0.4 mm diam., radially appressedly puberulous.
Infructescences subglobose, 3 – 4 cm diam., sometimes ± lobed, covered with c. 3 mm
long, cylindrical apices of the perianths; fruits ellipsoid, 1–1.5 cm long.
Distribution — Borneo.
Note — The material included in this species was initially referred to a subspecies
of A. melinoxylus which was included in the synonymy of A. chama Buch.-Ham. (see
Zhekun & Gilbert, Fl. China 5 (2003) 31), a species ranging from Sikkim to Vietnam or
part of a more broadly construed species, see note under A. hispidus. Artocarpus brevi
pedunculatus differs from A. chama (s.str.) in the cuneate to obtuse base of the lamina,
the absence of ramification of the lateral veins, the shorter stipules and peduncles.
5. Artocarpus elasticus Reinw. ex Blume

Artocarpus elasticus Reinw. ex [Blume, Cat. (1823) 101, nomen] Blume, Bijdr. (1825) 481; Miq., Fl.
Ned. Ind. 1, 2 (1859) 285; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211, p.p.; Koord. & Valeton,
Bijdr. Boomsoort. Java 11 (1906) 13; Koord., Exkurs.-Fl. Java 2 (1912) 92; W.H. Br., Min. Prod.
Philip. For. 2 (1921) 70, f. 24; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 251; Corner, Wayside
Trees Malaya (1940) 653, t. 193, 194; F.G. Browne, For. Trees Sar. & Brunei (1955) 352; F.M.
Jarrett, J. Arnold Arbor. 40 (1959) 346; Backer & Bakh.f., Fl. Java 2 (1965) 19; Kochummen, Tree
Fl. Malaya 3 (1978) 124, t. 3; Corner, Wayside Trees Malaya ed. 3, 2 (1988) 516, t. 157, 158, 159;
Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 196.
Artocarpus pubescens Blume, Bijdr. (1825) 481, non Willd. 1805; Moritzi, Syst. Verz. (1846) 75.
Artocarpus blumei Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 111, t. 4, f. 116; Miq. in Zoll., Syst. Verz.
2 (1854) 89; Fl. Ned. Ind. 1, 2 (1859) 285; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211; Kurz,
Natuurk. Tijdschr. Ned.-Indië 27 (1864) 182; Becc., For. Borneo (1902) 632.
Artocarpus kunstleri King in Hook.f., Fl. Brit. India 5 (1888) 540; Ann. Roy. Bot. Gard. (Calcutta) 2
(1889) 9, t. 4; Ridl., Trans. Linn. Soc. London, Bot. 3 (1893) 355; Fl. Malay Penins. 3 (1924) 353.
— Artocarpus blumei Trécul var. kunstleri (King) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 370.
Artocarpus scortechinii King in Hook.f., Fl. Brit. India 5 (1888) 542; Ann. Roy. Bot. Gard. (Calcutta)
2 (1889) 12, t. 9; Ridl., Fl. Malay Penins. 3 (1924) 355; Corner, Wayside Trees Malaya (1940)
657; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 344; Kochummen, Tree Fl. Malaya 3 (1978) 134, t. 6;
Corner, Wayside Trees Malaya ed. 3, 2 (1988) 521.
Artocarpus corneri Kochummen, Gard. Bull. Singapore 50 (1998) 197; Tree Fl. Sabah & Sarawak 3
(2000) 194, t. 3.
Artocarpus jarrettiae Kochummen, Gard. Bull. Singapore 50 (1998) 198; Tree Fl. Sabah & Sarawak
3 (2000) 198.
Tree up to 45(– 65) m tall, evergreen (or deciduous?), with buttresses. Leafy twigs
(5 –)7– 20 mm thick, brown to whitish appressedly puberulous to subhispidulous or to
hirtellous or hirsute, sometimes only on the scars of the stipules, ± scabrous or smooth,
drying dark brown to blackish; lenticels in the upper part of the internode; scars of the
stipules prominent. Leaves spirally arranged; lamina coriaceous to chartaceous, entire,
elliptic 13 – 40(– 60) by 6 – 20(– 35) cm, or when juvenile pinnately incised with 3 or
4 (or 5) pairs of lobes, apex short-acuminate, base rounded to cuneate (to subattenu-
ate), margin entire to repand to lobate (or denticulate towards the apex); upper surface
minutely whitish puberulous to (sub)hispidulous, on the main veins to hirtellous, ± sca
brous or smooth; lower surface brownish to whitish appressedly puberulous or partly
strigillose to (sub)hirtellous on the veins, scabridulous or smooth; lateral veins 10 –16
pairs, most of them branched or forked away from the margin, tertiary venation scalari
form, prominent; areoles usually bullate; petiole 2.5 –10 cm long, 2 – 6 mm thick, brown
appressedly puberulous, the epidermis persistent; stipules fully amplexicaul, 4 – 20 cm
long, brown (sub)hirsute to subvillous or to subsericeous, caducous. Staminate inflo
rescences axillary, solitary; peduncle 3.5 –7.5 or 9.5 –18 cm long, brown appressedly
puberulous; head cylindrical, 6 –15 by 1– 2.5 cm, sulcate, the grooves often interrupted
and almost straight; perianth tubular c. 0.6 mm long, the apex 2-lobed, minutely pu-
berulous; stamen c. 0.9 mm long, anther 0.2 – 0.4 mm long; interfloral bracts absent.
Pistillate inflorescences axillary, solitary; peduncle 4.5 –12 cm long, brown appressedly
puberulous; head ellipsoid to cylindrical; perianth tubular, brownish (to whitish) pu-
berulous to hispidulous, the apex convex to flat; stigma bifid (with equally or unequally
long arms); the flowers intermixed with subulate to filiform (± recurved) 6 –12 mm long
processes, these brown hispidulous or absent. Infructescences ellipsoid to cylindrical
(or to subglobose), (6 –)8 –12(–17) by 5.5(–10) cm, covered with 1– 4 mm long cushion
shaped to pyramidate to cylindrical apices of the perianths, intermixed with elongate
processes or not; fruits ellipsoid, 0.8 –1 cm long.
Distribution — Myanmar, Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo,
Java, Philippines (Palawan), Lesser Sunda Islands (Bali, Lombok, Sumbawa).
Habitat — Evergreen forest at low altitudes, sometimes up to c. 1500 m.
Uses — Cloth is made from bark. The infructescence has an unpleasant smell, but
the seed are eaten roasted.
Notes — 1. The pistillate inflorescences are quite different with regard to the pres-
ence of subulate to filiform processes among the flowers. The form without such proc-
esses has been recognised as Artocarpus scortechinii (Jarrett 1959). The upper surface
of the lamina of specimens with such inflorescences is often smooth, but can be sca
brous. The other form, in which the lamina is usually ± scabrous above, has elongated
processes and has been recognised as A. elasticus. The latter is more widespread than the
former, of which most collections (with a smooth upper surface of the lamina) have been
made in the Malay Peninsula and Sumatra, but some, including the type of A. corneri,
all with a scabrous upper surface of the lamina, are made in northern Borneo (Sarawak,
Brunei). As there are no other differentiating characters in the vegetative parts than the
inconsistent difference in the roughness of the upper surface of the lamina, and without
differences in the staminate inflorescences and flowers, distinction at the species level
is not justified. It might be convenient to indicate specimens (including the type of A.
jarrettiae) with interfloral processes as the ‘elasticus’-form, those without processes
and a smooth upper surface of the lamina as the ‘scortechinii’-form, and that without
processes but with the lamina ± scabrous above (from Borneo) as the ‘corneri’-form.
Artocarpus teijsmannii is another species with an even more erratic presence of
interfloral processes.
2. Artocarpus sericicarpus is very close to A. elasticus and included in the synonymy
of the latter by Kochummen (2000: 196). A form with interfloral processes, can be
distinguished by the ± patent brown hairs on the petiole and peduncle of the pistillate
inflorescence and is maintained as a distinct species, at least provisionally. Except for
the indumentum on the petiole and peduncle and the longer interfloral processes with
longer hairs, the two entities are similar. The similarities, the nature of the differences
and the slight overlap in the ranges of distribution provide arguments to reduce A.
sericicarpus to a subspecies. In Borneo (E Kutai) the two taxa have been collected in
the same locality, which is an indication that they are good species.
3. Artocarpus tamaran which shares the presence of elongate interfloral processes
and an uneven surface of the staminate inflorescence with A. sericicarpus and A. elas
ticus p.p. is distinct enough to be maintained as a distinct species.
4. The infructescences are eaten by monkeys; they are favoured by orang-utans.
6. Artocarpus excelsus F.M. Jarrett

Artocarpus excelsus F.M. Jarrett, Blumea 22 (1975) 409; [J. Arnold Arbor. 40 (1959) 340, as probably
new species]; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 197.
Tree up to 35 m tall, evergreen. Leafy twigs 3 – 4(– 5) mm thick, minutely whitish ap-
pressedly puberulous, smooth, drying blackish; lenticels scattered, few. Leaves spirally
arranged; lamina coriaceous, entire, elliptic, (4 –)6 –16 by (1.5 –)2 – 5 cm, apex shortly
(sub)acuminate, base cuneate to rounded, often slightly unequally, margin entire; upper
surface sparsely minutely whitish appressedly puberulous to (sub)glabrous, smooth;
lower surface minutely whitish appressedly puberulous on the midrib or also on the
lateral veins, smooth; lateral veins 14 – 20 pairs, tertiary venation scalariform, almost
flat; petiole (0.5 –)1– 2(– 3) cm long, 1– 2 mm thick, minutely whitish appressedly pu-
berulous, the epidermis persistent; stipules fully amplexicaul, 2 – 4.5 cm long, whitish
(to brownish) appressedly puberulous, on the whole outer surface or only on the keel,
caducous. Staminate inflorescences axillary, solitary; peduncle 1–1.5 cm long, minutely
whitish appressedly puberulous; head cylindrical to spicate, 1.5 – 20 by 0.5 cm; peri-
anth tubular c. 0.8 mm long, the apex 2-lobed, minutely puberulous; stamen c. 1 mm
long, anther c. 0.2 mm long; interfloral bracts absent. Pistillate inflorescences axillary,
solitary; peduncle 1.5 – 2.5 cm long, minutely whitish (appressedly) puberulous (or
also with patent brown uncinate hairs); head ellipsoid to cylindrical; perianth tubular,
sparsely minutely brownish puberulous, the apex convex; stigma simple; interfloral
bracts absent. Infructescences ellipsoid, up to 7 by 4 cm, covered with 1– 2 mm long
pyramidate apices of the perianths; fruits ellipsoid, c. 1.2 cm long.
Distribution — Borneo (Sabah, most collections from Mt Kinabalu).
Habitat — Evergreen forest, at altitudes between c. 1400 and 1800 m.
Note — This species is closely related to A. lowii, from which it mainly differs in
the smaller leaves; the status of subspecies should be considered.
7. Artocarpus heterophyllus Lam.

Artocarpus heterophyllus Lam., Encycl. 3 (1789) 210; Tabl. Encycl. (1797) t. 745; Trécul, Ann. Sci.
Nat., Bot. sér. 3, 8 (1847) 117; Miq., Fl. Ned. Ind. 1, 2 (1859) 287; Corner, Gard. Bull. Singapore
10 (1939) 56; Wayside Trees Malaya (1940) 654; Quisumb., Med. Pl. Philipp. (1951) 228; F.M.
Jarrett, J. Arnold Arbor. 40 (1959) 334; Backer & Bakh.f., Fl. Java 2 (1965) 19; Kochummen,
Tree Fl. Malaya 3 (1978) 127, t. 4; Corner, Wayside Trees Malaya ed. 3, 2 (1988) 517; Soepadmo,
Prosea 2 (1991) 86, cum t. — Artocarpus integrifolia L.f. var. heterophylla (Lam.) Pers., Syn. Pl.
2 (1807) 531.
Sitodium cauliflorum Gaertn., Fruct. Sem. Pl. 1 (1788) 345, t. 71, 72.
Artocarpus philippensis Lam., Encycl. 3 (1789) 210.
Polyphema jaca Lour., Fl. Cochinch. (1790) 546. — Artocarpus integrifolia L.f. var. glabra Stokes,
Bot. Mat. Med. 4 (1812) 331.
Artocarpus maxima Blanco, Fl. Filip. (1837) 669.
Artocarpus integrifolia auct. non L.f.: Willd., Sp. Pl. 4 (1805) 189; Pers., Syn. Pl. 2 (1807) 531; Roxb.,
Pl. Coromandel (1815) t. 250; Blume, Bijdr. (1825) 482; Hook., Bot. Mag. 55 (1828) t. 2883, 2884;
Roxb., Fl. Ind., ed. Carey 3 (1832) 522; Decne., Herb. Timorensis Descr. (1835) 169; Span., Lin-
naea 15 (1841) 343; Wight, Icon. Pl. Ind. Orient. 2 (1843) t. 678; Blanco, Fl. Filip. ed. 2 (1845)
466; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 115; Griff., Notul. Pl. Asiat. 4 (1854) 402; Miq.,
Fl. Ned. Ind. 1, 2 (1859) 287; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211; Blanco, Fl. Filip. ed.
3, 3 (1879) 75; Fern.-Vill., Nov. App. (1880) 203; King in Hook.f., Fl. Brit. India 5 (1888) 541;
Ann. Roy. Bot. Gard. (Calcutta). 2 (1889) 16 et errata (record from Deccan, India); Warb., Bot.
Jahrb. Syst. 13 (1891) 296; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 19; Elmer, Leafl.
Philipp. Bot. 2 (1909) 612; Koord., Exkurs.-Fl. Java 2 (1912) 93; Ridl., Fl. Malay Penins. 3 (1924)
351; Gagnep., Fl. Indo-Chine 5 (1928) 732; Subba Rao, J. Mysore Univ., B 1 (1940) 63.
Artocarpus integer auct. non (Thunb.) Merr.: Merr., Sp. Blancoan. (1918) 124; W.H. Br., Min. Prod.
Philip. For. 2 (1922) 262, t. 15, 16; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 41; Ochse & Bakh.,
Fruit (1931) 69, t. 28; Veg. Dutch East Indies (1931) 490; Burkill, Dict. Econ. Prod. Malay Penins.
(1935) 253; Merr., Comm. Fl. Cochinch. (1935) 135; Mendiola, Philip. Agric. 28 (1940) 789.
[Tsjaka-maram Rheede, Hort. Mal. 3 (1682) 17, t. 26 – 28.]
[Saccus arboreus major Rumph., Herb. Amboin. 1 (1741) 104, t. 30.]
Tree up to 10(– 30) m tall, sometimes with short buttresses, evergreen. Leafy twigs 2 –
6 mm thick, minutely whitish appressedly puberulous (to subpunctate), smooth, drying
brown to greyish; lenticels scattered. Leaves spirally arranged; lamina (sub)coriaceous,
entire (or if juvenile 3-lobed), elliptic to (sub)obovate, 4 –15 by 2 – 8(–12) cm, apex
short-acuminate to obtuse, base cuneate to rounded, mostly ± decurrent, margin entire,
± revolute; upper surface often initially with sparse (retrorse) whitish uncinate hairs,
soon glabrous, smooth; lower surface sparsely minutely whitish appressedly puberulous
on the main veins or only on the midrib, often initially also sparse (retrorse) uncinate
hairs, smooth ; lateral veins 6 –10 pairs, tertiary venation scalariform, slightly promi-
nent; petiole 0.8 – 3 cm long, 1– 2 mm thick, sparsely minutely whitish appressedly
puberulous, the epidermis persistent; stipules fully amplexicaul, 1– 5(– 8) cm long,
minutely whitish appressedly puberulous, caducous. Staminate inflorescences axillary
or cauliflorous on slender leafy branchlets, solitary; peduncle 1– 5.5 cm long, minutely
whitish appressedly puberulous, at the apex broadened into a rim; head cylindrical to
spicate or to subellipsoid, 2.5 –7 by 0.8 – 2.8 cm; perianth tubular 0.7–1 mm long, the
apex 2-lobed, minutely puberulous; stamen 1.5 – 2 mm long, anther c. 0.3 mm long;
flower intermixed with numerous (to few?) cylindrical processes, these as long as the
flowers, or processes absent. Pistillate inflorescences axillary, ramiflorous or cauli-
florous on leafy branchlets, solitary; peduncle 3 –10 cm long, minutely whitish (appress-
edly) puberulous, the apex broadened into a broad rim; head ellipsoidal to cylindrical;
perianth tubular, white puberulous to subhispidulous, the apex convex; stigma simple;
interfloral bracts (sometimes?) present, these peltate with the apical part 0.2 – 0.4 mm
diameter. Infructescences ellipsoid to narrowly ellipsoid to pyriform, 30 –100 by 25 – 50
cm, covered with c. 3 mm long conical to pyramidate apices of the perianths; fruits
ellipsoid, c. 3 cm long.
Distribution — In Malesia cultivated and (occasionally) naturalised in areas with
evergreen and semi-evergreen forest; probably (only) indigenous in western peninsular
India; cultivated throughout the tropics (under the names Jack, Jak, Jaquier, Jaca, etc.).
Uses — The sweet fleshy perianths in fruit are eaten; the seeds are eaten roasted or
boiled; the wood is used, e.g., for construction.
Notes — 1. There are two main forms in cultivation: one with firm fruiting perianths
and the other with soft ones.
2. The presence of processes among staminate flowers or bracts among pistillate
flowers is rather erratic.
8. Artocarpus hispidus F.M. Jarrett

Artocarpus hispidus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 149; Kochummen, Tree Fl. Malaya 3
(1978) 27, t. 4.
Tree up to 20 m tall, evergreen. Leafy twigs 4 –7 mm thick, ± densely brown hirsute,

the long hairs with swollen bases and intermixed with much shorter hairs, ± scabrous,
drying brown; lenticels scattered. Leaves spirally arranged; lamina subcoriaceous, en-
tire, elliptic to subobovate, (6 –)10 – 22 by (2.5 –)4.5 – 9 cm, short-acuminate to sub
acuminate or to acute (and apiculate) or to obtuse, base cuneate to obtuse, margin
dent(icul)ate towards the apex; upper surface brownish hispidulous, to hirtellous on
the main veins, ± scabrous; lower surface brownish (sub)hispidulous to puberulous, to
hirtellous on the main veins, scabridulous; lateral veins (8 –)12 –15 pairs, not branched
or forked away from the margin, tertiary venation scalariform (to reticulate), prominent;
petiole 1–1.5(– 2) cm long, 2 – 4 mm thick, brown hirtellous to subhirsute, the epider-

mis persistent; stipules 0.5 –1(– 2.5) cm, brown subhirtellous to strigose, caducous.
Staminate inflorescences axillary, solitary; peduncle 1– 3 cm long, brown hirtellous,
± scabrous; head ellipsoid to obovoid, 1.5 – 3 by 1– 2 cm; perianth tubular c. 0.6 mm
long, the apex 2-lobed, minutely puberulous; stamen c. 0.7 mm long, anther 0.1 mm
long; interfloral bracts peltate, the apical part 0.2 – 0.3 mm diam., yellowish puberulous.
Pistillate inflorescences axillary, solitary; peduncle 2.5 – 3.5 cm long, brown hirtellous,
± scabrous; head ellipsoid to subglobose; perianth tubular, hispidulous, the apex ±
convex; stigma simple; interfloral bracts peltate, the apical part 0.2 – 0.3 mm diam., yel-
lowish puberulous. Infructescences subglobose, 4 – 5.5 cm diam., covered with 5 – 6 mm
long cylindrical apices of the perianths; fruits ellipsoid, 1–1.3 cm long. — Fig. 18a.
Distribution — Malay Peninsula.
Habitat — Evergreen forest, at altitudes up to c. 350 m.
Notes — 1. This species resembles A. rigidus, from which it differs in the longer
peduncle of the staminate inflorescence, the denticulate margin of the lamina, and the
indumentum of the leafy twigs, consisting of long ± patent hairs with swollen bases,
intermixed with distinctly shorter hairs.
2. This species, A. asperulus Gagnep., A. calophyllus Kurz, A. hirsutus, and A. rigi
dus share the globose (to ellipsoid) infructescence densely covered with up to 3 – 9 mm
long cylindrical indurated apices of the flowers. The material north of the Malay Penin-
sula differs from A. hispidus and A. rigidus in the frequently branched or forked lateral
veins, the tendency of having fewer lateral veins, often up to 10 pairs, and a rounded to
6 cm
6 cm
a
2 cm
4 mm
f
1.5 mm
5 mm
1 mm
1 mm
c e d b h g 6 mm
i 2 mm
j
Fig. 18. a: Artocarpus hispidus F.M. Jarrett. Longitudinal section of infructescence. — b – i: Artocarpus
rigidus Blume. b. Leafy twig with pistillate inflorescence; c. pistillate inflorescence; d. young pistil-
late flower with interfloral bract; e. surface view of young pistillate inflorescence; f. surface view of
mature pistillate inflorescence; g. longitudinal section of portion of mature pistillate inflorescence;
h. tangential section of portion of mature pistillate inflorescence; i. longitudinal section of staminate
inflorescence; j. surface view of staminate inflorescence (a: Corner SFN 37035; b, d–h, i, j: 24 T 1P
195; c: 24 T 27).
subcordate base of the lamina. Artocarpus hirsutus from the Indian Peninsula is distinct
in the pendulous spicate staminate inflorescences and the absence of interfloral bracts in
the pistillate inflorescence; the other species have subglobose staminate inflorescences
as in A. hispidus and A. rigidus. In the material occurring north of Peninsular Malaysia,
the indumentum on leafy twigs and petioles can be appressed or ± patent, the stipules
are of widely varying length, the margin of the lamina is entire or denticulate, the up-
per surface of the lamina smooth or ± scabrous, and the length of the peduncles varies
considerably. Some of this variation may just enter Peninsular Malaysia, in Kedah and
Langkawi Island (in the material indicated by Kochummen 1978: 132) as the ‘asperu-
lus’-form. Pending the results of further study on the complex in continental Asia and
its relation to the Malesian region, it is preferable to recognise for Malesia A. hispidus
and A. rigidus, both as species and not as forms as has been proposed by Kochummen
(1978), where the sparsely hairy and more densely hairy forms of A. rigidus are indi-
cated as var. ‘glabra’ and var. ‘tomentosa’, respectively.
3. The suggested study should also include A. chama Buch.-Ham. (= A. chaplasha
Roxb. = A. melinoxylus Gagnep.) which in vegetative characters largely matches the
continental material circumscribed above, differing only in the shorter apices of the
pistillate flowers which are up to 4 cm long (or up to 6 mm long according to Gagnep.,
Fl. Indo-Chine 5 (1928) 734).
9. Artocarpus integer (Thunb.) Merr.

Artocarpus integer (Thunb.) Merr., Interpr. Herb. Amboin. (1917) 190; Corner, Gard. Bull. Singapore
10 (1939) 56, t. 1, 2; Wayside Trees Malaya ed.1 (1940) 655; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
329; Backer & Bakh.f., Fl. Java 2 (1965) 19; Kochummen, Tree Fl. Malaya 3 (1978) 127; Corner,
Wayside Trees Malaya ed. 3, 2 (1988) 518, t. 157; P. C.M. Jansen, Prosea 2 (1991) 91, cum t.;
Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 199. — Radermachia integra Thunb., Kongl.
Svenska Vetensk. Acad. Handl. 37 (1776) 254; Houtt., Nat. Hist. II, 11 (1779) 453. — Sitodium
macrocarpon Thunb., Philos. Trans. 69 (1779) 467, substitute name. — Artocarpus macrocarpon
(Thunb.) Dancer, Cat. Bot. Gard. Jamaica 1 (1792); Mabb., Taxon 30 (1981) 12. — Artocarpus
integrifolia L.f., Suppl. (1781) 411, substitute name. — Artocarpus integrifolia L.f. var. hirsuta
Stokes, Bot. Mat. Med. 4 (1812) 330.
Artocarpus jaca Lam. var. ß, Encycl. 3 (1789) 209.
Polyphema champeden Lour., Fl. Cochinch. (1790) 546, pro max. parte. — Artocarpus champeden
(Lour.) Stokes, Bot. Mat. Med. 4 (1812) 330; Spreng., Syst. Veg. 3 (1826) 804; Merr., Interpr.
Herb. Amboin. (1917) 190; Ochse & Bakh., Fruit (1931) 67, t. 27; Veg. Dutch East Indies (1931)
486; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 249; Merr., Comm. Fl. Cochinch. (1935) 135;
Mendiola, Philip. Agric. 28 (1940) 789.
Artocarpus polyphema Pers., Syn. 2 (1807) 531; Blume, Bijdr. (1825) 481; Span., Linnaea 15 (1841)
343; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 115; Miq., Fl. Ned. Ind. 1, 2 (1859) 286; Hoola
van Nooten, Fl. Fr. & Feuill. Java (1863) t. 26; Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211;
King in Hook.f., Fl. Brit. India 5 (1888) 542; Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 11, t. 7B;
Warb., Bot. Jahrb. Syst. 13 (1891) 296; Ridl., Trans. Linn. Soc. London, Bot. 3 (1893) 355; Koord.
& Valeton, Bijdr. Boomsoort. Java 11 (1906) 21; Koord., Exkurs.-Fl. Java 2 (1912) 94; Ridl., Fl.
Malay Penins. 3 (1924) 354; Gagnep., Fl. Indo-Chine 5 (1928) 734.
Artocarpus hirsutissima Kurz, Natuurk. Tijdschr. Ned.-Indië 27 (1864) 182.
Artocarpus integer (Thunb.) Merr. var. silvestris Corner, Gard. Bull. Singapore 10 (1939) 76; F.M.
Jarrett, J. Arnold Arbor. 40 (1959) 351; Kochummen, Tree Fl. Malaya 3 (1978) 128, t. 4.
[Saccus arboreus minor Rumph., Herb. Amboin. 1 (1741) 107, t. 31.]
Tree up to c. 30 m tall, evergreen. Leafy twigs 2 – 4 mm thick, whitish puberulous

(or punctate) and also brown hirtellous to (sub)hirsute, sometimes only (and sparsely)
on and near the nodes, the hairs with swollen bases and sometimes uncinate, some-
times glabrous, drying dark brown to blackish; lenticels scattered, few. Leaves spi-
rally arranged; lamina subcoriaceous, entire (or when juvenile to 3-lobate), elliptic to
(sub)obovate, (4 –)8 – 20(– 27) by (1.5 –)4 – 8(–12) cm, apex acuminate, base cuneate
to obtuse to rounded, often slightly unequally, margin entire, ± revolute; upper surface
whitish puberulous and brownish strigillose on the whole surface or only on the main
veins or midrib, or glabrous; lower surface whitish puberulous and brown hirtellous
on the whole surface or only on the main veins, the hairs all or partly uncinate, or
(becoming) glabrous, smooth; lateral veins 6 –12 pairs, tertiary venation towards the
margin (obliquely) scalariform or largely reticulate, slightly prominent to almost flat;
petiole 0.8 – 3 cm long, 1– 2 mm thick, brown (sub) hirsute or also whitish puberulous,
the epidermis persistent; stipules fully amplexicaul, 1.5 – 5(– 9) cm long, (sparsely)
brown (sub)hirsute to hirtellous to strigillose, the hairs with ± swollen bases and some
uncinate, or also whitish puberulous, caducous. Staminate inflorescences axillary or
cauliflorous on slender leafy branchlets, solitary; peduncle 2.5 – 6 cm long, brown hirtel-
lous, also white puberulous; head narrowly ellipsoid to cylindrical to spicate, 1.5 – 5.5
by 0.9 –1.2 cm; perianth tubular 0.7–1 mm long, the apex 2-lobed, minutely puberulous;
stamen 1–1.3 mm long, anther 0.2 – 0.3 mm long; interfloral bracts absent. Pistillate
inflorescences axillary, ramiflorous or cauliflorous on slender leafy branchlets, solitary;
peduncle 1.5 – 6.5(–10) cm long, brown hirtellous to (retrorsely) strigillose and most
hairs uncinate or also whitish puberulous; head cylindrical to subobovoid; perianth tu-
bular, brown puberulous to subhispidulous, the apex truncate to convex; stigma simple;
interfloral bracts absent. Infructescences narrowly ellipsoid to ellipsoid to ovoid (to
subglobose), 15 – 35 by 10 –15 cm, covered with 1.5 – 3 mm long, cylindrical to conical
to pyramidate apices of the perianths; fruits ellipsoid, c. 3 cm long.
Distribution — Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo, Java,
Celebes, Moluccas (Sula Islands, Ambon), New Guinea (western).
Notes — 1. Several collections from the Malay Peninsula and Sumatra have been
included in var. silvestris, distinct by the odourless and almost tasteless infructescences
as well as by the colour of withering leaves (see Jarrett 1959). The material tends to
be glabrous or glabrescent (losing uncinate hairs on the lamina, mainly on the midrib
beneath and the margin). Corner (1939: 75) regarded the variety as the wild form of the
species.
2. As explained by Corner in much detail (1939: 56 – 81), Radermachia integra is
based on different elements. The type sheet comprises material with hirsute leafy twigs
of the present species together with material without such indumentum which belongs
to A. heterophyllus, the Jack. This has created considerable (nomenclatural) confusion,
partly because of moving in renaming (as Sitodium macrocarpon) and in describing the
accent from the first element to the second (Thunberg, Kongl. Svenska Vetensk. Acad.
Handl. 37 (1779)). The element on the type sheet that should function as the type for
A. integer was clearly indicated by (Svedelius and) Corner (1939: plates 1 and 2).
10. Artocarpus kemando Miq.

Artocarpus kemando Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 418; Becc., For. Borneo (1902) 628;
Ridl., Fl. Malay Penins. 5 (1925) 335; Merr., Pl. Elmer. Born. (1929) 46; Burkill, Dict. Econ. Prod.
Malay Penins. (1935) 256; Corner, Gard. Bull. Singapore 10 (1939) 282; Wayside Trees Malaya
(1940) 656; F.G. Browne, For. Trees Sar. & Brunei (1955) 353; F.M. Jarrett, J. Arnold Arbor. 40
(1959) 354; Kochummen, Tree Fl. Malaya 3 (1978) 129, t. 5; Corner, Wayside Trees Malaya ed. 3,
2 (1988) 519; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 200.
Artocarpus maingayi King in Hook.f., Fl. Brit. India 5 (1888) 542; Ann. Roy. Bot. Gard. (Calcutta) 2
(1889) 11, t. 8A; Ridl., Fl. Malay Penins. 3 (1924) 354; Corner, Gard. Bull. Singapore 10 (1939)
282; Wayside Trees Malaya (1940) 657; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 357; Kochummen,
Tree Fl. Malaya 3 (1978) 131, t. 5; Corner, Wayside Trees Malaya ed. 3, 2 (1988) 520.
Artocarpus brunneifolia S. Moore, J. Bot. 63, Suppl. (1925) 112.
Artocarpus sumatranus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 353.
Tree up to 35 m tall, evergreen, with (short) buttresses. Leafy twigs 2 – 3 mm thick,

brown (to whitish) appressedly puberulous to appressed-hispidulous or to hirtellous
(with part of the hairs uncinate and ± retrorse), scabridulous, drying (dark) brown;
lenticels scattered. Leaves spirally arranged; lamina coriaceous, entire, elliptic to sub
obovate, (3 –)7–15(–18) by (1.5 –)3 –10 cm, apex short-acuminate to obtuse or to round-
ed, base cuneate to rounded, margin entire, ± revolute; upper surface (sparsely) minutely
whitish puberulous, on the midrib the hairs often longer (and brownish), or subglabrous,
smooth; lower surface (sparsely) minutely whitish puberulous, on the main veins the
hairs longer and brownish, smooth; lateral veins 8 –15 pairs, usually unbranched, ter-
tiary venation scalariform, slightly prominent; petiole 1– 3 cm long, 1– 2 mm thick,
brownish appressedly puberulous, the epidermis persistent; stipules fully amplexicaul,
0.7– 2 cm long, brown appressedly puberulous to strigillose, caducous. Staminate inflo
6 cm
2.5 cm
a
c
b
d
6 cm
1 cm
Fig. 19. Artocarpus kemando Miq. a. Leafy twig with pistillate inflorescence; b. leaf; c. pistillate
inflorescence; d. staminate inflorescence (a – c: no. 90 E I O 755; d: FRI bb 19850).
rescences axillary, solitary (or in pairs) peduncle (0.3 –)0.7–1.5(– 2) cm long, brown
velutinous or appressedly puberulous; head cylindrical to spicate, 2 – 4 by 0.3 – 0.5 cm,
(irregularly) sulcate, occasionally subtended by some 0.5 –1 mm long scale-like bracts;
perianth tubular c. 0.4 mm long, the apex 2-lobed, minutely puberulous; stamen 0.4 – 0.5
mm long, anther c. 0.2 mm long; interfloral bracts absent. Pistillate inflorescences axil-
lary, solitary (or in pairs); peduncle 1– 4 or 0.3 – 0.8 cm long, brown velutinous; head
ellipsoid to ovoid, occasionally subtended by some 0.5 –1 mm long scale-like bracts;
perianth tubular, densely brown puberulous, the apex flat to concave; stigma simple;
interfloral bracts absent. Infructescences ellipsoid to subglobose, up to 4 by 2.5 cm,
covered with c. 2 mm long (concave) cushion-shaped (or pyramidate to 3 mm long)
apices of the perianths; fruits ellipsoid, c. 0.8 cm long. — Fig. 19.
Distribution — Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo.
Habitat — Evergreen forest at low altitudes, often in swampy places.
Notes — 1. The differences between A. kemando and A. maingayi as recognised
by Jarrett (1959) are so small that uniting them seems to be justified. Material ranked
under the latter always has peduncles shorter than 1 cm and the lamina tends to have a
rounded apex.
2. The differences between A. kemando and A. sumatranus are not of such importance
that recognition of the latter as species is justified.
11. Artocarpus lanceifolius Roxb.

Artocarpus lanceifolius Roxb. [Hort. Bengal. (1814) 103, nomen] Fl. Ind., ed. Carey 3 (1832) 527, as
‘lanceofolia’ and ‘lanceaefolia’, respectively; Wight, Icon. Pl. Ind. Orient. 2 (1843) t. 679; Trécul,
Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 122; King in Hook.f., Fl Brit. Ind. 5 (1888) 543; Ann. Roy.
Bot. Gard. (Calcutta) 2 (1889) 13, t. 11; Renner, Bot. Jahrb. Syst. 39 (1907) 366; Ridl., Fl. Malay
Penins. 3 (1924) 354; Foxworthy, Malayan Forest Rec. 3 (1927) 128, with plates; Burkill, Dict.
Econ. Prod. Malay Penins. (1935) 256; Corner, Wayside Trees Malaya (1940) 656, t. 197; F.M.
Jarrett, J. Arnold Arbor. 40 (1959) 140; Kochummen, Tree Fl. Malaya 3 (1978) 129, t. 5; Corner,
Wayside Trees Malaya ed. 3, 2 (1988) 519, t. 157, 160; Kochummen, Tree Fl. Sabah & Sarawak
3 (2000) 2001.
Artocarpus clementis Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 164. — Artocarpus lancei
folius Roxb. subsp. clementis (Merr.) F.M. Jarrett, J. Arnold Arbor. 40 (1959) 142.
Tree up to 35 m tall, evergreen. Leafy twigs 6 – 8 mm thick, ± densely brown (to

whitish) appressedly puberulous, drying brown; lenticels scattered. Leaves spirally
arranged; lamina coriaceous, entire, elliptic, sometimes 3-lobed to pinnately incised,
10 – 40 by 3.5 –17(– 30) cm, apex short-acuminate, base cuneate to obtuse, margin
entire to repand (to sublobate); upper surface glabrous, smooth; lower surface gla-
brous or sparsely whitish appressedly puberulous on the midrib, smooth; lateral veins
9 –14 pairs, mostly some of them forked away from the margin or branched venation
scalariform, slightly prominent; petiole 1– 3(– 5.5) cm long, 2 – 5 mm thick, minutely
whitish appressedly puberulous, the epidermis persistent; stipules fully amplexicaul,
1– 4.5 cm long, brown appressedly puberulous, caducous. Staminate inflorescences
axillary, solitary; peduncle 2 –7 cm long, brown (to whitish) appressedly puberulous;
head ellipsoid to subobovoid to narrowly elliptic to cylindrical, 3 – 6 by 1–1.5(– 2) cm;
perianth tubular c. 2 mm long, the apex 2-lobed, minutely puberulous to subglabrous;
stamen 3 – 3.5 mm long, anther c. 0.3 mm long; interfloral bracts peltate, the apical part
0.3 – 0.4 mm diam., radially appressedly puberulous. Pistillate inflorescences axillary,
solitary; peduncle 2.5 –7 cm long, densely brown (to whitish) appressedly puberulous;
head subglobose; perianth tubular, brown puberulous, the apex ± convex; stigma bifid;
interfloral bracts peltate, caducous or persistent, the apical part 0.3 – 0.4 mm diam.,
radially appressedly puberulous. Infructescences subglobose, 6 –7 cm diam., sometimes
± lobed, covered with 3 – 4 mm long, cylindrical apices of the perianths; fruits ellipsoid,
1–1.5 cm long.
Distribution — Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo (north-
eastern).
Habitat — Evergreen forest at altitudes up to c. 1200 m.
Uses — The pulp around seeds is eaten by man.
Note — The material from Borneo, recognised by Jarrett (1959) as a distinct sub-
species, differs slightly from that from Thailand, Sumatra, and the Malay Peninsula,
in the usually more slender petioles and persistent interfloral bracts of the pistillate
inflorescence.
12. Artocarpus lowii King

Artocarpus lowii King in Hook.f., Fl. Brit. India 5 (1888) 542; Ann. Roy. Bot. Gard. (Calcutta) 2 (1889)
10, t. 7A; Ridl., Fl. Malay Penins. 3 (1924) 353; Burkill, Dict. Econ. Prod. Malay Penins. (1935)
257; Corner, Gard. Bull. Singapore 10 (1939) 283; Wayside Trees Malaya (1940) 656; F.M. Jarrett,
J. Arnold Arbor. 40 (1959) 339; Kochummen, Tree Fl. Malaya 3 (1978) 131; Corner, Wayside Trees
Malaya ed. 3, 2 (1988) 519.
Tree up to 25 m tall, evergreen. Leafy twigs 3 – 6 mm thick, minutely whitish ap-

pressedly puberulous, smooth, drying blackish; lenticels scattered, few. Leaves spirally
arranged; lamina (sub)coriaceous, entire, narrowly elliptic, 10 – 36(–55) by 4 –13(– 20)
cm, apex short-acuminate to acute, base cuneate (to obtuse), often slightly unequally,
margin entire to repand; upper surface (sub)glabrous, smooth; lower surface minutely
whitish appressedly puberulous on the midrib, smooth; lateral veins 11–16 pairs, terti-
ary venation scalariform, slightly prominent; petiole 1.5 – 3.5 cm long, 1–2 mm thick,
minutely whitish appressedly puberulous or also with sparse brown uncinate hairs, the
epidermis persistent; stipules fully amplexicaul, 1– 5(– 8) cm long, minutely whitish
appressedly puberulous, caducous. Staminate inflorescences axillary, solitary; peduncle
c. 4 cm long, minutely whitish appressedly puberulous or also with brown (retrorse)
uncinate hairs; head cylindrical to spicate or to subellipsoid, 2 – 2.5 by c. 0.5 cm; peri-
anth tubular c. 0.8 mm long, the apex 2-lobed, minutely puberulous; stamen c. 1.5 mm
long, anther c. 0.2 mm long; interfloral bracts absent. Pistillate inflorescences axillary,
solitary; peduncle 3.5 – 5.5 cm long, minutely whitish (appressedly) puberulous or also
with brown (retrorse) uncinate hairs; head ellipsoid to cylindrical; perianth tubular,
minutely brownish puberulous, the apex convex; stigma simple; interfloral bracts ab-
sent. Infructescences cylindrical, up to 6.5 by 3.5 cm, covered with 1– 2 mm long,
pyramidate apices of the perianths; fruits ellipsoid, c. 0.7 (?) cm long.
Distribution — Sumatra (eastern), Malay Peninsula.
Uses — The milk sap is greasy and used as ointment.
13. Artocarpus obtusus F.M. Jarrett

Artocarpus obtusus F.M. Jarrett, Blumea 22 (1975) 410; Kochummen, Tree Fl. Sabah & Sarawak 3
(2000) 204.
Tree up to 35 m tall, base of trunk fluted, evergreen. Leafy twigs 3 – 6 mm thick,

minutely whitish appressedly puberulous, smooth, drying blackish; lenticels scattered,
few. Leaves spirally arranged; lamina coriaceous, entire, elliptic, 18 – 23 by 4 –19 cm,
apex rounded to obtuse, base rounded to obtuse (to subcordate), often slightly unequally,
margin entire, ± revolute; both upper and lower surface (sub)glabrous and smooth;
lateral veins 7–10 pairs, some of them branched or forked away from the margin, terti-
ary venation scalariform, flat; petiole 1– 3.5 cm long, 2 – 3 mm thick, (sub)glabrous,
the epidermis persistent; stipules fully amplexicaul, 1– 4 cm long, minutely brownish
to whitish appressedly puberulous, caducous. Staminate inflorescences axillary, soli-
tary; peduncle 1.2 – 3.5 cm long, sparsely minutely whitish puberulous and also with
brown uncinate hairs; head ovoid to conical, up to 2.5 by 1 cm; perianth tubular, c. 0.7
mm long, the apex 2-lobed, minutely puberulous; stamens immature; interfloral bracts
absent. Pistillate inflorescences axillary, solitary; peduncle 3 –7.5 cm long, minutely
whitish puberulous and also with brown uncinate hairs; head ellipsoid; perianth tubular,
brownish hispidulous, the apex convex; stigma bifid; flowers intermixed with interflo-
ral processes. Infructescences ellipsoid, up to 8 by 6 cm, covered with 2 – 3 mm long,
pyramidate apices of the perianths, intermixed with 6 –10 mm long spine-like, slightly
curved processes; fruits ellipsoid to subglobose, c. 1.5 cm long.
Distribution — Borneo (Sarawak).
Note — The species is characterised by (sub)glabrous lamina with rounded apex.
14. Artocarpus odoratissimus Blanco

Artocarpus odoratissimus Blanco, Fl. Filip. (1837) 671; Fern.-Vill., Nov. App. (1880) 203; Elmer,
Leafl. Philipp. Bot. 2 (1909) 618; Wester, Philipp. Agric. Rev. 8 (1915) 108, t. 7d, 9b; Merr.,
J. Straits Branch Roy. Asiat. Soc. 76 (1917) 80; Sp. Blancoan. (1918) 124; Enum. Philipp. Flow.
Pl. 2 (1923) 42; Wester, Philipp. Agric. Rev. 17 (1924) 24; Bull. Bur. Agric. Philipp. 39 (1924) 129,
t. 15b, 32b; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 147; F.S. dela Cruz, Prosea 2 (1991) 94, cum t.;
Artocarpus tarap Becc., For. Borneo (1902) 626.
Artocarpus mutabilis Becc., For. Borneo (1902) 627.
Tree up to 25 m tall, evergreen. Leafy twigs 4 –12 mm thick, ± densely brown strig-
illose to hispidulous, ± scabrous or hirsute, the long hairs with swollen bases, inter-
mixed with much shorter hairs or not, drying brown; lenticels scattered. Leaves spirally
arranged; lamina subcoriaceous (to chartaceous), entire, elliptic (to subrhombic) to
obovate, 10 – 30(– 55) by (6 –)10 –15(– 30) cm, apex subacuminate to apiculate, base
cuneate to subdecurrent, margin entire, repand, denticulate, or (unilaterally) lobate
(when juvenile 3-lobed to pinnately incised with 1 or 2 pairs of lateral lobes); upper
surface brownish hispidulous, on the main veins to strigillose, or hirtellous, ± scabrous;
lower surface brownish (sub)hispidulous to puberulous, on the main veins to strigil-
lose, or hirtellous, scabridulous; lateral veins 13 – 20 pairs, often branched or forked
away from the margin, tertiary venation scalariform, prominent; petiole (1.5 –)2 – 3 cm
long, 3 – 4 mm thick, brown strigillose (to hirtellous), or hirsute, the epidermis persist-
ent; stipules 1– 5(– 8) cm, brown strigillose to subhirtellous or to strigose, or hirsute,
caducous. Staminate inflorescences axillary, solitary; peduncle (2.5 –)3 –13 cm long,
brown appressedly puberulous to hispidulous or hirtellous to subhirsute, ± scabrous;
head ellipsoid to obovoid to clavate, 4 – 9 by 2.5 – 3.5 cm; perianth tubular c. 1.8 mm
long, the apex 2-lobed, minutely puberulous; stamen c. 2 mm long, anther 0.1– 0.2
mm long; interfloral bracts peltate, the apical part 0.3 – 0.4 mm diam., yellowish pu-
berulous. Pistillate inflorescences axillary, solitary; peduncle 5 –14 cm long, densely
brown appressedly puberulous to hispidulous, or hirtellous to subhirsute, ± scabrous;
head ellipsoid to subglobose; perianth tubular, hispidulous or hispid, the apex ± convex;
stigma simple; interfloral bracts peltate, the apical part 0.3 – 0.4 mm diam., yellowish
puberulous. Infructescences subglobose, 8 –15 cm diam., sometimes ± lobed, fruit-
containing part 5.5 –12 cm diam., covered with 8 –12 mm long cylindrical apices of the
perianths; fruits ellipsoid, c. 1 cm long.
Distribution — Borneo, Philippines (Mindoro, Mindanao, Basilan, Sulu Archipela-
go; see note 2).
Notes — 1. Two forms can be recognised: one with short ± appressed hairs and the
other with long ± patent hairs with swollen bases, hirsute on the leafy twigs, petioles,
and stipules and hirtellous on the lamina and the hairs on the pistillate perianth longer.
Only one of the collections examined, W. Meijer 1851, from Tarakan, is more or less
intermediate. The latter form is indicated on some labels as subsp. barbatum by Jarrett,
an unpublished name.
2. According to Jarrett (1959) the species might have been introduced to the Philip-
pines from Borneo.
15. Artocarpus rigidus Blume

Artocarpus rigidus Blume, Bijdr. (1825) 482; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 114; Miq.
in Zoll., Syst. Verz. 2 (1854) 89, 95; Miq., Fl. Ned. Ind. 1, 2 (1859) 286; Fl. Ned. Ind., Eerste
Bijv. (1861) 418; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211; King in Hook.f., Fl. Brit. India 5
(1888) 540; Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 8, t. 3; Koord. & Valeton, Bijdr. Boomsoort.
Java 11 (1906) 17; Koord., Exkurs.-Fl. Java 2 (1912) 93; Ridl., Fl. Malay Penins. 3 (1924) 352;
K. Heyne, Nutt. Pl. Ned.-Ind. (1927) 564; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 258; Cor-
ner, Wayside Trees Malaya (1940) 657, t. 198, 199; F.G. Browne, For. Trees Sar. & Brunei (1955)
353; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 150; Kochummen, Tree Fl. Malaya 3 (1978) 131, t. 6;
Corner, Wayside Trees Malaya ed. 3, 2 (1988) 521, t. 157, 163, 164; Kochummen, Tree Fl. Sabah
& Sarawak 3 (2000) 208, t. 5; B. Seibert & P.C.M. Jansen, Prosea 2 (1991) 81, cum t.
Artocarpus echinata Roxb., [Hort. Bengal. (1814) 66, nomen] Fl. Ind., ed. Carey 3 (1832) 527; Wight,
Icon Pl. Ind. Orient. 2 (1843) t. 680; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 113.
Artocarpus cuspidatus Griff., Notul. Pl. Asiat. 4 (1854) 400.
Artocarpus kertau Zoll. ex Miq. in Zoll., Syst. Verz. 2 (1854) 89, 95; Miq., Fl. Ned. Ind. 1, 2 (1859)
287; Koord., Exkurs.-Fl. Java 2 (1912) 95.
Artocarpus dimorphophylla Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 417.
Artocarpus varians Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 417.
Artocarpus chaplasha auct. non Roxb.: Gagnep., Fl. Indo-Chine 5 (1928) 735.
Tree up to 35 m tall, with buttresses, evergreen. Leafy twigs (1–)2 – 8 mm thick,

± densely brown strigillose, scabridulous, drying brown; lenticels scattered. Leaves
spirally arranged; lamina (sub)coriaceous, entire, elliptic to (sub)obovate, (3 –)10 –

20(– 30) by (1.5 –)5 –10(–15) cm (or pinnately lobed to parted and up to 55 by 35 cm),
rounded to short-(sub)acuminate to apiculate, base obtuse to cuneate or to rounded,
margin entire to repand (or crenate towards the apex), ± revolute; upper surface brown-
ish puberulous to densely strigillose or to only (sparsely) strigillose on the midrib,
smooth or scabridulous; lower surface brownish strigillose on the main veins to pu-
berulous or to hispidulous on the smaller veins, smooth or ± scabrous; lateral veins
(6 –)10 – 20 pairs, none (or rarely one) branched or forked away from the margin, ter-
tiary venation scalariform, ± prominent; petiole (0.5 –)1– 2(– 2.5) cm long, 2 – 3 mm
thick, brown strigillose, the epidermis persistent; stipules (0.5 –)1– 2(– 4) cm, brown
strigillose to strigose, caducous. Staminate inflorescences axillary, solitary; peduncle
0.2 –1 cm long, brownish strigillose, ± scabrous; head obovoid to subglobose, 1.2 – 2 cm
diam.; perianth tubular, c. 1.2 mm long, the apex 2-lobed, minutely puberulous; stamen
c. 1.3 mm long, anther 0.1 mm long; interfloral bracts peltate, the apical part 0.2 – 0.3
mm diam., yellowish puberulous. Pistillate inflorescences axillary, solitary; peduncle
0.8 – 3(– 4) cm long, brown strigillose, ± scabrous; head ellipsoid to subglobose (to
depressed-globose); perianth tubular, hispidulous to muriculate, the apex ± convex;
stigma simple; interfloral bracts peltate, the apical part 0.2 – 0.3 mm diam., yellowish
puberulous. Infructescences subglobose, 4 – 9 cm diam., covered with 3 – 8 mm long,
cylindrical apices of the perianths; fruits ellipsoid, 1.2 –1.5 cm long. — Fig. 18b – i.
Distribution — Myanmar, Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo,
Java, Lesser Sunda Islands (Bali?).
Habitat — Evergreen forest at altitudes up to 700(–1000) m.
Note — This species is closely related to A. hispidus, under which the differences
and relations to material and names in continental Asia are discussed.
16. Artocarpus sarawakensis F.M. Jarrett

Artocarpus sarawakensis F.M. Jarrett, Blumea 22 (1975) 410; Kochummen, Tree Fl. Sabah & Sarawak
3 (2000) 210.
Tree up to 25 m tall, evergreen. Leafy twigs 4 –7 mm thick, ± densely brown hirsute

(to substrigose), long hairs with a swollen base intermixed with minute whitish hairs
or else densely brown puberulous, drying (dark) brown; lenticels scattered. Leaves spi-
rally arranged; lamina coriaceous, entire, elliptic to subobovate, 25 – 45 by 14 – 25 cm,
apex short-acuminate, base cuneate (to obtuse), margin repand; upper surface sparsely
brown hirtellous on the midrib, glabrescent, smooth; lower surface whitish to brownish
strigillose to appressedly puberulous on the main veins or also sparsely brown subhir-
sute on the midrib, scabridulous; lateral veins 12 – 20 pairs, some forked away from
the margin, tertiary venation scalariform, slightly prominent; petiole 1– 2.5 cm long,
2 – 5 mm thick, brown hirsute, the long hairs intermixed with minute whitish hairs, the
epidermis persistent; stipules fully amplexicaul, 1– 4(– 6) cm long, brown to yellowish
hirsute or hirtellous, caducous. Staminate inflorescences axillary, solitary; peduncle
3.5 – 8 cm long, brown hirsute to strigose, the long hairs intermixed with minute whitish
hairs; head obovoid to subobovoid to cylindrical, 3 – 6 by 1– 2.5 cm; perianth tubular
c. 0.8 mm long, the apex 2-lobed, minutely puberulous to subglabrous; stamen c. 1.3
mm long, anther c. 0.1 mm long; interfloral bracts peltate, the apical part c. 0.2 mm
diam., puberulous. Pistillate inflorescences: peduncle c. 5.5 cm long, brown hirsute to
strigose, the long hairs intermixed with minute whitish hairs or brown subvelutinous and
with retrorse uncinate hairs; head subglobose; perianth tubular, brown puberulous, the
apex ± convex; stigma bifid; interfloral bracts peltate, the apical part c. 0.2 or c. 5 mm
diam., puberulous (with hairs of similar or different length). Infructescences subglobose
or ellipsoid, 3 – 5 cm diam., covered with cushion-shaped 2 – 3 mm long apices of the
perianths; fruits ellipsoid, c. 1.3 cm long.
Distribution — Sumatra (Riau Province), Borneo (Sarawak).
Notes — 1. This species shows similarities to A. lanceifolius, particularly the large-
leaved specimens from the Malay Peninsula, but differs in the presence of hairs on the
lamina below and the ± patent (longer) hairs on the stipules.
2. The species also resembles A. odoratissimus from which it differs in the coriaceous
lamina, smooth above and with a repand margin.
3. The single collection from Sumatra differs from the (five examined) collections
from Sarawak in the absence of long hairs on the leafy twig, petiole, and midrib of the
lamina lower surface. The hairs on the stipules are shorter and the dense short brown
indumentum of the peduncle is intermixed with longer retrorse uncinate hairs. The api-
cal parts of the peltate interfloral bracts are larger.
17. Artocarpus sepicanus Diels

Artocarpus sepicanus Diels, Bot. Jahrb. Syst. 67 (1935) 176; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
362.
Tree up to 40 m tall, evergreen, sometimes with buttresses; bark flaking off. Leafy
twigs 3 – 5 mm thick, brown, glabrous, drying dark brown to blackish; lenticels scat-
tered. Leaves spirally arranged; lamina coriaceous, entire, elliptic to subobovate,
(6 –)10 – 25(– 30) by (2.5 –)5 –10(–12) cm, apex short-acuminate, base subcordate to
rounded, unequal, margin entire to repand; both surfaces glabrous, smooth; lateral veins
9 –16 pairs, tertiary venation scalariform, almost flat; petiole 1– 2.5 cm long, 1– 2 mm
thick, glabrous, the epidermis flaking off; stipules fully amplexicaul, 1– 3 cm long,
glabrous, caducous. Staminate inflorescences axillary, solitary; peduncle 1.5 – 3 cm
long, glabrous; head cylindrical to spicate, 2 – 3 by 0.5 – 0.7 cm; perianth tubular c. 0.8
mm long, the apex 2-fid, minutely puberulous; stamen c. 1 mm long, anther c. 0.4 mm
long; interfloral bracts peltate, the apical part 0.4 – 0.5(–1) mm diam., ciliolate. Pistillate
inflorescences axillary, solitary; peduncle 2.5 – 4.5 cm long, glabrous; head cylindrical;
perianth tubular, minutely puberulous, the apex convex; stigma bifid; interfloral bracts
peltate, the apical part 0.4 – 0.5(–1) mm diam., ciliolate. Infructescences ellipsoid to
subglobose, up to 4.5 by 1.5 cm, covered with 1– 2 mm long cushion shaped apices of
the perianths; fruits ellipsoid, c. 0.5 cm long.
Note — This species is characterised by the exfoliating epidermis of the petiole and
the glabrous vegetative parts.
18. Artocarpus sericicarpus F.M. Jarrett

Artocarpus sericicarpus F.M. Jarrett, J. Arnold Arbor. 40 (1959) 350; Kochummen Tree Fl. Sabah &
Sarawak 3 (2000) 196.
Artocarpus elasticus auct. non Blume: Fern.-Vill., Nov. App. (1880) 202; Stapf, Kew Bull. (1894)
108; Wester, Philipp. Agric. Rev. 8 (1915) 109, t. 8a; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 41;
Wester, Bull. Bur. Agric. Philipp. 39 (1924) 78, t. 196, 32 c; W.H. Br., Useful Pl. Philipp. (1941)
463, f. 188.
Artocarpus blumei auct. non Trécul: S. Vidal, Revis. Pl. Vasc. Filip. (1886) 254; Elmer, Leafl. Philipp.
Bot. 2 (1909) 613.
Tree up to 40 m tall, evergreen. Leafy twigs 10 – 20 mm thick, minutely brown to

whitish appressedly puberulous and also brown strig(ill)ose to subhirsute, smooth,
drying dark brown to blackish; lenticels in the upper part of the internode; scars of the
stipules prominent. Leaves spirally arranged; lamina subcoriaceous, entire, elliptic,
20 – 40(–70) by 10 – 25(– 50) cm, or when juvenile ± pinnately incised with 2 or 3 pairs
of lobes, apex short-acuminate, base rounded to cuneate, margin entire to repand or
lobate; upper surface sparsely brown hirtellous on the midrib, smooth; lower surface
brownish puberulous to strigillose on the smaller veins to subhirsute on the main veins
or midrib, usually scabridulous; lateral veins 10 –16 pairs, most of them branched or
forked away from the margin, tertiary venation scalariform, prominent; areoles usually
bullate; petiole 2 –7.5 cm long, 3 – 6 mm thick, minutely brownish appressedly puberu-
lous and also brown hirtellous to subhirsute or strigillose, the epidermis persistent;
stipules fully amplexicaul, 6 –12 cm long, brown subhirsute to subvillous, caducous.
Staminate inflorescences axillary, solitary; peduncle 3 –10 cm long, brown appressedly
puberulous; head cylindrical, 3.5 –10 by 1.5 – 2 cm, sulcate, the grooves ± (spirally)
twisted; perianth tubular c. 1.5 mm long, the apex 2-lobed, minutely puberulous; stamen
c. 1.8 mm long, anther c. 0.4 mm long; interfloral bracts absent. Pistillate inflorescences
axillary, solitary; peduncle 6 –18 cm long, brown appressedly puberulous and hirtel-
lous; head ellipsoid to cylindrical; perianth tubular, brownish appressedly puberulous,
the apex convex; stigma simple, twisted; the flowers intermixed with 1– 3.5 cm long
filiform processes, these brown puberulous to strigillose hirtellous and thickened at the
apex. Infructescences ellipsoid to cylindrical or to subglobose, up to (6 –)8 –12(–17)
by 5.5(–10) cm, covered with 3 – 4 mm long conical apices of the perianths (hidden by
the processes); fruits ellipsoid, c. 1 cm long.
Distribution — Borneo, Philippines (Luzon, Samar, Biliran, Negros, Mindanao),
Celebes, Moluccas (Talaud and Sula Islands, Buru).
Habitat — Evergreen forest at low altitudes, sometimes up to c. 1000 m.
Note — This species is closely related to A. elastica, as discussed under the latter.
19. Artocarpus tamaran Becc.

Artocarpus tamaran Becc., For. Borneo (1902) 626; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 352;
Tree up to 40 m tall, evergreen, with buttresses. Leafy twigs 5 –10 mm thick, whit-
ish puberulous and brown hirsute to subvillous, smooth, drying dark brown to black-
ish; lenticels scattered. Leaves spirally arranged; lamina coriaceous, entire, elliptic to
narrowly elliptic to subovate, 20 – 35 by 11–17 cm, or when juvenile, up to c. 100 cm

long, pinnately incised (down to the midrib), with up to 8 pairs of segments, the ‘rachis’
often narrowly winged, apex short-acuminate to subacute, base rounded to obtuse,
margin entire to repand, ± revolute; upper surface (sub)glabrous, smooth; lower sur-
face whitish puberulous on the veins and brown strig(ill)ose to hirtellous on the main
veins, smooth; lateral veins (13 –)18 – 24 pairs, rarely branched or forked away from
the margin, slightly impressed above, tertiary venation scalariform, prominent; petiole
3.5 – 4 cm long, 2 – 3 mm thick, whitish puberulous and brown subhirsute, the epidermis
persistent; stipules fully amplexicaul, 3 – 9 cm long, (dark) brown (sub)hirsute to subvil-
lous, caducous (or subpersistent). Staminate inflorescences axillary, solitary; peduncle
3.5 – 5.5 cm long, brown appressedly puberulous; head cylindrical, 6 –7 by 1–1.5 cm,
tuberculate with 2 – 3 mm high cushion shaped processes covered by dark brown hairs;
perianth tubular c. 0.6 mm long, the apex 2-lobed, minutely puberulous; anther c. 0.2
mm long; interfloral bracts absent. Pistillate inflorescences axillary, solitary; pedun-
cle 5.5 –10 cm long, whitish puberulous and brown hirsute; head ellipsoid; perianth
tubular brown puberulous to hispidulous, the apex convex; stigma simple; the flowers
intermixed with 6 –10 mm long filiform processes, these brown retrorsely puberulous
to hispidulous. Infructescences ellipsoid, up to 10(–14) by 5(– 8) cm, covered with 3 – 4
mm long conical to cushion shaped apices of the perianths, intermixed with elongate
processes; fruits ellipsoid, c. 0.6 cm long.
Uses — Cloth is made from bark.
Note — This species differs from A. elasticus and A. sericicarpus, e.g., in the more
numerous lateral veins which are usually not branched or forked and the tuberculate
surface of the staminate inflorescence.
20. Artocarpus teijsmannii Miq.

Artocarpus teijsmannii Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 418; Boerl., Handl. Fl. Ned. Ind. 3
(1900) 333, 371; K. Heyne, Nutt. Pl. Ned.-Ind. (1927) 565; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
341; Kochummen, Tree Fl. Malaya 3 (1978) 134, t. 6; Tree Fl. Sabah & Sarawak 3 (2000) 211.
Artocarpus peduncularis Kurz, J. Bot. 13 (1875) 331; King in Hook.f., Fl. Brit. India 5 (1888) 541;
Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 10, t. 6; Becc., For. Borneo (1902) 632; Ridl., Fl. Malay
Penins. 3 (1924) 353; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 258; Corner, Gard. Bull.
Singapore 10 (1939) 283; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 341.
Tree up to 40 m tall, sometimes with buttresses, evergreen. Leafy twigs 3 – 8 mm

thick, minutely whitish to brownish appressedly puberulous or glabrous, smooth, dry-
ing blackish to brown or to greyish; lenticels (very) few, scattered; scars of the stip-
ules prominent. Leaves spirally arranged; lamina (sub)coriaceous, entire, elliptic to
(sub)ovate, (5 –)10 – 25(– 40) by (3 –)6 –13 (–17) cm, apex short-acuminate to acute or
to obtuse, base cuneate to rounded (to subcordate), often slightly unequal, margin entire,
often ± revolute; upper surface minutely whitish puberulous, the hairs slightly longer
and appressed on the main veins, or glabrous, smooth; lower surface minutely whit-
ish puberulous, the hairs slightly longer and appressed on the main veins, or glabrous,
smooth; lateral veins 6 –16(–18) pairs, often branched or forked away from the margin,
tertiary venation scalariform, almost flat; petiole (1–)2 – 5 cm long, (1–)2 – 3 mm thick,
minutely whitish appressedly puberulous or glabrous, the epidermis persistent; stipules
fully amplexicaul, 1– 5(– 8) cm long, whitish to brownish appressedly puberulous, to-
wards the apex brownish subsericeous, or glabrous, caducous. Staminate inflorescences
axillary, solitary; peduncle 2 – 8 cm long, minutely whitish appressedly puberulous or
subglabrous; head spicate, 3.5 –7.5 by 0.5 – 0.7(– 0.9) cm; perianth tubular c. 1.5 mm
long, the apex 2-lobed, minutely puberulous; stamen 1– 2 mm long, anther c. 0.5 mm
long; flowers intermixed with 1– 2 mm long subulate processes or with peltate bracts,
the apical part 0.2 – 0.3(–1) mm diameter. Pistillate inflorescences axillary, solitary;
peduncle 3 –10 cm long, minutely whitish (appressedly) puberulous or subglabrous;
head cylindrical; perianth tubular, minutely brownish to whitish puberulous, the apex
convex; stigma simple or bifid; flowers often intermixed with 0.3 – 0.6 mm long subulate
processes. Infructescences cylindrical to clavate, up to 8.5 by 4.5 cm, covered with 1– 2
mm long and pyramidate, or sometimes up to 6 mm long and subulate, apices of the
perianths, (in some infructescences none, in others some or numerous) up to 6 mm long
subulate processes; fruits ellipsoid, c. 0.7 (?) cm long.
Note — The species can be subdivided into two disjunct subspecies.
a. subsp. teijsmannii
Artocarpus peduncularis Kurz, J. Bot. 13 (1875) 331; King in Hook.f., Fl. Brit. India 5 (1888) 541;
Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 10, t. 6; Becc., For. Borneo (1902) 632; Ridl., Fl. Malay
Penins. 3 (1924) 353; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 258; Corner, Gard. Bull.
Singapore 10 (1939) 283; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 341.
Leafy twigs, leaves, and stipules whitish to brownish minutely appressedly puberu-
lous; stipules brown subsericeous or with a comose apex. Peduncles minutely puberu-
lous. Perianth of the pistillate flower puberulous.
Distribution — India (Nicobar Islands), Thailand; in Malesia: Sumatra, Malay Pe-
ninsula, Borneo, Celebes, Moluccas (Ceram, Sula Islands), New Guinea (western).
Habitat — Evergreen forest at low altitudes, often in swampy areas.
Notes — 1. The minute hairs on both the upper and lower surface of the lamina and
the brown subsericeous or comose apex of the stipules are characteristic for this sub
species.
2. Subulate processes (1– 2 mm in length) are present in staminate inflorescences
of some collections, but in those made in New Guinea the flowers are intermixed
with peltate bracts. The occurrence of subulate interfloral processes is rather erratic in
the pistillate inflorescences: they vary from totally absent to few scattered among the
flowers or concentrated at the base of the head, or else may occur abundantly. The proc-
esses may be abundant on one or two inflorescences on a branch and totally absent on
the others. The situation can be confusing if the apices of the perianths also become
subulate. These subulate structures can be distinguished from the processes by the
perforation at the top through which the style or stigma is protruding.
3. In most pistillate inflorescences the stigmas are simple, but some may be bifid. In
others, such as Moll BW 11572, all stigmas are bifid.
b. subsp. subglabrus C.C. Berg

Artocarpus teijsmannii Miq. subsp. subglabrus C.C. Berg, Blumea 50 (2005) 543.
Ficus ralumensis K. Schum., Notizbl. Bot. Gart. Berlin-Dahlem 2 (1898) 112; Fl. Schutzgeb. Südsee
(1901) 282; Diels, Bot. Jahrb. Syst. 67 (1935) 231; Corner, Gard. Bull. Singapore 21 (1965) 99;
C.C. Berg, Flora Malesiana, Ser. I, 17-2 (2005) 702.
Leafy twigs, leaves, and stipules glabrous. Peduncles glabrous or with few uncinate
hairs. Perianth of the pistillate flower puberulous only at the apex.
Distribution — Solomon Islands (Guadalcanal); in Malesia: New Guinea (Bismarck
Archipelago, Morobe Province).
Note — Because only two fertile collections are known, it is not certain whether
the variation in the presence of interfloral bracts and processes as described for subsp.
teijsmannii applies to this subspecies; the only collection with a staminate inflorescence
has interfloral bracts, of which those at the base of the head are large, c. 1 mm diam.
and the pistillate inflorescences does not have processes among the pyramidate perianth
apices. Both simple (in the collection from the Solomon Islands) and bifid stigmas (in
New Guinea) are found.
21. Artocarpus treculianus Elmer

Artocarpus treculianus Elmer, Leafl. Philipp. Bot. 2 (1909) 617; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 43; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 302.
Artocarpus nigrescens Elmer, Leafl. Philipp. Bot. 2 (1909) 614; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 42.
Artocarpus ovatifolia Merr., Philipp. J. Sci., Bot. 9 (1914) 268; Enum. Philipp. Flow. Pl. 2 (1923) 42.
Artocarpus ovatifolia Merr. var. dolichostachys Merr., Enum. Philipp. Flow. Pl. 2 (1923) 43.
Artocarpus communis auct. non J.R. & G. Forst.: Merr., Philipp. J. Sci., Bot. 3 (1908) 401.
Tree up to 15(– 40?) m tall, with small buttresses, evergreen. Leafy twigs 4 –10 mm
thick, whitish minutely appressedly puberulous to strigillose, drying dark brown to
blackish; lenticels scattered. Leaves spirally arranged; lamina (sub)coriaceous, entire,
elliptic to subrhombic, 15 – 28 by 9 –17 cm, or pinnatifid, with 1– 4 pairs of lateral lobes,
18 – 28 by 11– 24 cm, base cuneate to rounded, apex subacuminate to acute, margin
subentire to repand; upper surface white appressedly puberulous on the main veins or
only at the base of the midrib, sometimes also sparsely hirtellous on the midrib, smooth;
lower surface whitish puberulous on the main veins, scabridulous; lateral veins 10 –13
pairs, at least the lower ones mostly branched or forked away from the margin, tertiary
venation scalariform, almost flat; petiole (2 –)3 – 8 cm long, 1.5 – 3 mm thick, whitish
appressedly puberulous, the long hairs intermixed with much shorter hairs; stipules
6 –18 cm long, whitish appressedly puberulous, caducous. Staminate inflorescences
axillary, solitary; peduncle 1.2 – 2.5 cm long, whitish appressedly puberulous; head
spicate, 1–21 cm long, 0.4 – 0.7 cm diam.; perianth tubular, 1–1.8 mm long, the apex
2-lobed, minutely puberulous; stamen 1.5 – 2.8 mm long, anther 0.3 – 0.5 mm long;
interfloral bracts absent. Pistillate inflorescences axillary, solitary; peduncle 1.5 – 5 cm
long, whitish appressedly puberulous; head ellipsoid; perianth tubular brown to yellow-
ish puberulous to muriculate, the apex convex; stigma bifid; interfloral bracts absent.
Infructescences ellipsoid, up to 7 by 5 cm, often irregularly shaped, covered with c. 3
mm long cylindrical apices of the perianths; fruits ellipsoid, c. 1.2 cm long.
Distribution — Philippines (widespread).
Habitat — (Semi-)evergreen forest at altitudes up to c. 800 m.
Notes — 1. Jarrett (1959: 305) described several staminate collections deviating in
larger anthers (0.6 – 0.8 mm long) and/or the presence of interfloral subulate processes
(indicated as sterile flowers); these anomalies have not been found in the material ex-
amined for the present treatment. The discrepancies may be caused by mixed collections
as the size of the anthers matches that of A. altilis which also has interfloral bracts or
processes, see note under A. altilis (p. 86).
2. Collection Elmer 12468 differs from the other collections by lacking the branched
lateral veins.
3. The majority of the collections have entire laminas but some have pinnately in-
cised ones which resemble those of A. altilis and apparently also occur in adult speci-
mens.
4. This species shows similarities to A. teijsmannii and is probably more closely
related to the latter than to A. altilis, as suggested by Jarrett (1959).
Subgenus Pseudojaca
Artocarpus J. & G. Forst. subg. Pseudojaca Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 117; F.M. Jar-
rett, J. Arnold Arbor. 41 (1960) 73. — Artocarpus J. & G. Forst. sect. Pseudojaca (Trécul) Renner,
Bot. Jahrb. Syst. 39 (1907) 368.
Artocarpus J. & G. Forst. subg. Pseudojaca Trécul sect. Pseudojaca (Trécul) Renner ser. Peltati F.M.
Jarrett, J. Arnold Arbor. 41 (1960) 83.
Artocarpus J. & G. Forst. subg. Pseudojaca Trécul sect. Pseudojaca (Trécul) Renner ser. Clavati F.M.
Jarrett, J. Arnold Arbor. 41 (1960) 130.
Artocarpus J. & G. Forst. subg. Pseudojaca Trécul sect. Glandulifolium F.M. Jarrett, J. Arnold Arbor.
41 (1960) 134.
Leaves alternate and distichous, entire (when adult); lamina with glands on the mar-
gin, on the midrib above the basal lateral veins, or in their axils and the axils of other
lateral veins; stipules lateral, short, 0.3 – 0.5(–1) cm long. Inflorescences axillary (or
staminate ones on short-shoots on the older wood), interfloral bracts peltate (or cylin-
drical or spathulate). Staminate flowers with 2 – 4, partly connate (or free) tepals. Free
parts of apices of pistillate flowers mostly not prominent, in some species (sometimes)
forming low pyramidate hardened processes.
DISTRIBUTION
The subgenus is represented by up to 21 species in an area ranging from Sri Lanka

to the Solomon Islands and Australia, mostly as elements of evergreen forest. The
subgenus is concentrated in the western Malesian region and the adjacent part of the
Asian mainland. The most widespread species is A. lacucha, from India and China to
the Solomon Islands. Fourteen species are found in the Malesian region, the others in
China and Indochina. Four of them have been treated by Jarrett (1960), four from China
(Sichuan and Yunnan) were added (see Zhekun & Gilbert, Fl. China 5 (2003) 32 – 33),
and one is described from Thailand (see below).
MORPHOLOGY
Habit — Several species show intermittent growth with differences in the colour and
indumentum of the current growth compared to that of the previous season. In some
species the transition is marked by a concentration of the stipular scars which formed
terminal buds. In some species, as A. tomentosulus, lateral buds are also formed. In-
termittent growth is associated with deciduousness, at least in A. lacucha. The lamina
of species with intermittent growth is subcoriaceous to chartaceous and the margin is
sometimes (in A. lacucha nearly always) denticulate towards the apex or (only) at the
acumen.
Indumentum — Uncinate hairs are often present, especially in the juvenile state. They
can be consistently present on some parts of the plant and may then have diagnostic
value.
The following groups of species can be recognised based on the presence, colour,
and denseness of the indumentum of twigs and laminas.
1) Hairs present in the areoles, and not only on the venation: A. fulvicortex, A. glaucus
(in Malesia and Australia), A. hypargyreus Hance (China), A. styracifolius Pierre
(China and Indochina), and A. thailandicus C.C. Berg (Blumea 50 (2005) 531 from
northern Thailand, and probably also two or three of the more recently described
Chinese species (see above).
2) Hairs predominantly brown(ish) and usually dense on various parts: A. albobrun
neus (?), A. fulvicortex, A. lacucha, A. longifolius, A. subrotundifolius, and A. to
mentosulus. This group has intermittent growth (and can be deciduous).
3) Hairs predominantly whitish and sparse on leafy twigs and leaves (except for the
hairy form of A. gomezianus in India and Sri Lanka): A. gomezianus, A. nitidus,
A. reticulatus, A. rubrovenius, and A. vrieseanus. This group probably consists of
evergreen species without intermittent growth.
Leaves — The leaves are occasionally opposite (in A. lacucha).
Lamina — The base of the lamina is sometimes ‘peltate’ with a narrow strip of laminar
tissue connecting both sides of the base over the apex of the petiole. This phenomenon is
found in several species. The base of the lamina is often ± unequal, probably correlated
with the distichous arrangement of the leaves.
Glands — The glandular-crenate margin of the lamina of A. altissimus is highly pecu-
liar, as such glands are not known in other Moraceae. Almost as peculiar are the (waxy?)
glandular spots on the midrib above the base and in the axils of the lateral veins of the
lamina lower surface. The glands on the midrib can be short or long, extending to the
next lateral vein. They are found in all species with the exception of A. altissimus, but
are not always manifest; more in the species with hairy leaves than in those with sub-
glabrous leaves. They are in their positions quite similar to those in numerous Ficus
species.
Stipules — The stipules are mostly 0.3 – 0.5 cm long, in some species on opening shoots
up to 1 cm long, and then often subpersistent. Stipules cover terminal and lateral (rest-
ing) buds.
Bracts — Interfloral bracts are mostly peltate with short stalks. They are (nearly) al-
ways caducous though sometimes persistent in pistillate inflorescences of some species;
shedding is not a consistent character. In A. styracifolius Pierre (China and Indochina)
the interfloral bracts are represented by cylindrical processes up to c. 5 mm long and
± curved in pistillate inflorescences (in fruit). The bracts of staminate inflorescences of
A. petelotii Gagnep. (China and Indochina) are spathulate (according to Jarrett 1960:
130).
In several species, the bracts at the base of the head are basally attached instead of
peltate and somewhat larger than the others.
Pistillate flowers — The apices of pistillate flowers usually form (hairy) rims which
surround the emerging stigma and persist in fruit. In some species the apices become
low and hardened pyramidate processes: always in A. reticulatus and sometimes in
A. lacucha.
DELIMITATION AND SUBDIVISION
This subgenus clearly differs from subg. Artocarpus in the distichous arrangement of
the leaves, the short(er than 1 cm) non-amplexicaul (lateral) stipules, the presence of
glands and glandular spots, and the mostly smooth surface of the infructescence.
The three groups of species which can be recognised by their indumentum and the
highly peculiar A. altissimus (sect. Glandulifolium F.M. Jarrett) could be regarded as
natural subdivisions. It is doubtful whether the separation of species into a group with
peltate interfloral bracts (ser. Peltati) and a group with non-peltate ones (ser. Clavati)
reflects natural alliances since modified peltate bracts (cylindrical processes) also occur
in subg. Artocarpus and in the African genus Treculia
KEY TO THE SPECIES OF SUBGENUS PSEUDOJACA
1a. Margin of lamina with glands. — Sumatra, Borneo . . . . . . . . . . . 2. A. altissimus

b. Margin of lamina without glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Lower surface of lamina hairy in the areoles, ± glaucous when fresh . . . . . . . . . . 3
b. Lower surface of lamina hairy only on the venation . . . . . . . . . . . . . . . . . . . . . . . 4
3a. Indumentum on veins of lower surface of lamina whitish. — Sumatra, Malay
Peninsula, Borneo, Java, Lesser Sunda Islands . . . . . . . . . . . . . . . . . 4. A. glaucus
b. Indumentum on veins of lower surface of lamina brown. — Sumatra, Malay Penin-
sula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. A. fulvicortex
4a. Lamina ± densely hairy on midrib above and venation beneath . . . . . . . . . . . . . 5
b. Lamina sparsely hairy to subglabrous on midrib above and venation beneath
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
5a. Petiole 0.2 – 0.8 cm long and (2 –)3 – 4 mm thick; indumentum brown. — Borneo
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. A. longifolius
b. Petiole at least 0.8 cm long, or if shorter, then c. 1.5 cm thick and the indumentum
whitish (and occurring in New Guinea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Lateral veins 6 –10 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Lateral veins 10 –20 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7a. Apex of the lamina rounded, leafy twigs for the greater part sparsely hairy, petiole
0.8 –1.5 cm long. — Borneo . . . . . . . . . . . . . . . . . . . . . . . . 1. A. albobrunneus
b. Apex of the lamina acuminate, or if rounded, then the leafy twigs densely hairy
and the petiole usually 3 –7 cm long. — Widespread . . . . . . . . . . . 6. A. lacucha
8a. Lateral veins for the greater part furcate far from the margin or branched. — Phil-
ippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. A. subrotundifolius
b. Lateral veins not or sometimes one or few furcate far from the margin . . . . . . . 9
9a. Epidermis of petiole flaking off; margin of lamina entire. — Borneo . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. A. tomentosulus
b. Epidermis of petiole (usually) persistent; margin of the lamina (usually) denticu-
late towards the apex, at least at the acumen. — Widespread . . . . 6. A. lacucha
10a. Lamina ± scabrous beneath; base of lamina often minutely auriculate. — Celebes,
Moluccas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. A. reticulatus
b. Lamina (usually) smooth beneath; base of lamina never auriculate . . . . . . . . . 11
11a. Lateral veins 5 –10(–11) pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
b. Lateral veins (8 –)10 – 20 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12a. Head of staminate inflorescence ellipsoid to obovoid, 1–1.5(– 2) cm diam. and
infructescence to c. 8 cm diameter. — Philippines . . . . . . . . 10. A. rubrovenius
b. Head of staminate inflorescence subglobose, or if ellipsoid to obovoid, then 0.2 –
0.5 cm diam. and infructescence to c. 4.5 cm diameter. — Philippines . . . . . . 13
13a. Peduncle of staminate inflorescence 1.2 –1.7 cm long, peduncle of pistillate inflo-
rescence (in fruit) 2.5 – 5.5 cm long. — Celebes, Moluccas . . . 13. A. vrieseanus
b. Peduncle of staminate inflorescence 0.1– 0.4 cm long; peduncle of pistillate inflo-
rescence (in fruit) 0.1– 0.5(– 2.5) cm long. — Sumatra, Malay Peninsula, Borneo,
Philippines, New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. A. nitidus
14a. Peduncle of staminate inflorescence 0.5 –1.7 cm long and peduncle of pistillate
inflorescence (in fruit) 1.5 – 4.5 cm long. — Sumatra, Malay Peninsula, Java,
Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. A. gomezianus
b. Peduncle of staminate inflorescence 0.1– 4 cm long and peduncle of pistillate
inflorescence (in fruit) usually 0.1– 0.5 cm long (or in Borneo up to 2.5 cm long,
but then the epidermis of the petiole flaking off instead of persistent). — Sumatra,
Malay Peninsula, Borneo, Philippines, New Guinea . . . . . . . . . . . . 8. A. nitidus
1. Artocarpus albobrunneus C.C. Berg

Artocarpus albobrunneus C.C. Berg, Blumea 50 (2005) 541.
Tree, evergreen (?). Leafy twigs 2 – 3 mm thick, sparsely brown minutely puberu-
lous, more densely hairy towards the base of the petiole, smooth, drying dark brown
to blackish. Leaves distichous; lamina coriaceous, elliptic to subobovate, 6 –13 by
2.5 – 6.5 cm, apex rounded, base rounded to cordulate, slightly unequal, margin entire,
± revolute (towards the base); upper surface brownish minutely puberulous mainly on
the midrib, smooth; lower surface minutely brownish puberulous on the main veins to
whitish puberulous on the smaller veins, smooth; lateral veins 7– 9(–10) pairs, none
of them forked away from the margin, tertiary venation scalariform, (and also the
smaller veins) prominent; petiole 0.8 –1.5 cm long, 1–1.5 mm thick, densely minutely
brown puberulous, the epidermis flaking off; stipules lateral, 0.2 – 0.4 cm long, brown-
ish puberulous, caducous. Staminate inflorescences not seen. Pistillate inflorescences
axillary, solitary; (in fruit) peduncle 1.5 – 2.5 cm long, densely brownish puberulous;
head subglobose (?); stigma simple; interfloral bracts peltate, the apical part c. 0.2 mm
diam., puberulous, persistent in the central part of the infructescence, largely caducous
in the peripheral part. Infructescences discoid and strongly lobed around the fruits, the
central part whitish velutinous, the peripheral part (lobes) red-brown velutinous.
Distribution — Borneo (E Kalimantan).
Note — This species is unusual among the species of Artocarpus subg. Pseudojaca
in the coriaceous lamina that dries greenish. The different colour of parts of the infruc
tescence are curious, – a way to create contrast colours to attract dispersers?
2. Artocarpus altissimus (Miq.) J.J. Sm.

Artocarpus altissimus (Miq.) J.J. Sm., Icon. Bogor. 3 (1907) 79, t. 233; [Teijsm. & Binn., Cat. Hort.
Bot. Bogor. (1866) 85, nomen]; Douglas & Baas-Beck., Bull. Jard. Bot. Buitenzorg III, 17 (1947)
197, 291; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 135. — Morus ? altissima Miq., Fl. Ned. Ind.,
Eerste Bijv. (1861) 415.
Grewia ? subcordata Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 404; Burret, Notizbl. Bot. Gart. Berlin-
Dahlem 9 (1926) 736.
Tree up to 30 m tall, sometimes with buttresses, deciduous. Leafy twigs 1.5 – 3 mm

thick, brownish to whitish puberulous, sometimes partly with retrorse uncinate hairs,
smooth, drying dark brown to blackish. Leaves distichous; lamina subcoriaceous to
chartaceous, entire or 3 – 5-lobate when juvenile, ovate to subovate to elliptic to narrow-
ly elliptic, 6 – 20 by 5 –13 cm, apex acuminate, base cordate to subcordate to rounded,
(almost) equal, margin glandular-crenate, ± revolute, at least towards the base; upper
surface whitish to brownish puberulous on the main veins (to subglabrous), smooth
(to scabridulous); lower surface sparsely to densely whitish to brownish puberulous to
subtomentose on the (main) veins, smooth (to scabridulous); lateral veins 5 – 9 pairs, the
basal pair branched, tertiary venation scalariform, ± prominent; petiole 1– 3.5 cm long,
1.5 – 2.5 mm thick, ± densely brownish to whitish puberulous, the epidermis flaking off;
stipules lateral, 0.3 – 0.5 cm long, brownish to whitish appressedly puberulous, cadu-
cous. Staminate inflorescences axillary or on short-shoots on the older wood; peduncle
5 –7 mm long, brownish velutinous; head ellipsoid to cylindrical, 5 – 8 mm long, 2 – 3
mm diam.; perianth with 4 free tepals, c. 0.7 mm long; stamen not exserted, anther c.
0.4 mm long; interfloral bracts peltate, the apical part c. 0.6 mm diam., puberulous.
Pistillate inflorescences axillary or on short-shoots on the older wood; peduncle up
to 12 mm long, brownish velutinous; head globose, at anthesis c. 5 mm diam.; stigma
bifid; interfloral bracts peltate. Infructescences not seen.
Distribution — Thailand; in Malesia: Sumatra, Borneo (western).

Notes — 1. This rare species is distinguishable by the glandular margin of the
lamina.
2. Fertile herbarium material is scarce and, therefore, the species description is in-
complete.
3. Artocarpus fulvicortex F.M. Jarrett

Artocarpus fulvicortex F.M. Jarrett, J. Arnold Arbor. 41 (1960) 116; Corner, Wayside Trees Malaya
(1940) 658, t. 197, ‘Artocarpus sp.’; Kochummen, Tree Fl. Malaya 3 (1978) 126, t. 4; Corner,
Wayside Trees Malaya ed. 3, 2 (1988) 517, t. 157, 160.
Tree up to 35 m tall, with intermittent growth and terminal and axillary buds, decidu-
ous. Leafy twigs 2.5 – 6 mm thick, densely brown puberulous, smooth, drying brown
to blackish. Leaves distichous; lamina (sub)coriaceous, entire, elliptic to subrotundate,
9 – 21 by 5.5 –16 cm, apex short-acuminate to rounded, base rounded to cuneate (to sub-
attenuate), ± unequal, margin entire, revolute to flat; upper surface brownish puberulous
mostly on the main veins, smooth; lower surface brownish strigillose to appressedly
puberulous on the midrib, patently puberulous on the smaller veins, minutely whitish
appressedly puberulous in the areoles, smooth, nearly glabrous when fresh; lateral
veins 6 –10 pairs, often one or more forked away from the margin, tertiary venation
scalariform, prominent, also the smaller veins; petiole (1.5 –)3 –7 cm long, (1.5 –)2.5 – 3
mm thick, densely brown puberulous, the epidermis persistent; stipules lateral, 0.3 – 0.5
cm long, brownish to yellowish puberulous, caducous. Staminate inflorescences axil-
lary, solitary (or in pairs); peduncle 1– 2 mm long, brown short-velutinous; head sub
globose, 4 – 6 mm diam.; perianth with 2 partly connate tepals, c. 0.7 mm long; stamen
c. 0.8 mm long, anther c. 0.2 mm long; interfloral bracts peltate, the apical part 0.3 – 0.5
mm diam., puberulous. Pistillate inflorescences axillary, solitary (or in pairs); peduncle
2 – 4 mm long, brown short-velutinous; head subglobose; stigma simple; interfloral
bracts peltate, persistent, the apical part 0.3 – 0.5 mm diam., puberulous, persistent.
Infructescences subglobose, up to 4 cm diam., subvelutinous, smooth to scabridulous;
fruits ellipsoid, c. 1 cm long.
Distribution — Sumatra, Malay Peninsula.
4. Artocarpus glaucus Blume

Artocarpus glaucus Blume, Bijdr. (1825) 483; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 121; Miq.
in Zoll., Syst. Verz. 2 (1854) 90, 96; Miq., Pl. Jungh. (1854) 293; Fl. Ned. Ind. 1, 2 (1859) 288;
Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 212; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906)
26; Koord., Exkurs.-Fl. Java 2 (1912) 95; 4 (1924) 479; F.G. Browne, For. Trees Sar. & Brunei
(1955) 353; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 118; Backer & Bakh.f., Fl. Java 2 (1965) 19;
Kochummen, Tree Fl. Malaya 3 (1978) 126; Chew, Fl. Australia 3 (1989) 23; Kochummen, Tree
Fl. Sabah & Sarawak 3 (2000) 198.
Artocarpus glaucescens Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 120; Miq. in Zoll., Syst. Verz. 2
(1854) 90; Fl. Ned. Ind. 1, 2 (1859) 288; Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 212.
Artocarpus zollingeriana Miq. in Zoll., Syst. Verz. 2 (1854) 90, 95; Fl. Ned. Ind. 1, 2 (1859) 289.
Artocarpus biformis Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 419.
Artocarpus tephrophylla Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 422. — Artocarpus glaucescens Trécul
var. tephrophylla (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 212.
Artocarpus denisoniana King in Hook.f., Fl. Brit. India 5 (1888) 544; Ann. Roy. Bot. Gard. (Calcutta)
2 (1889) 14, t. 8B; Ridl., Fl. Malay Penins. 3 (1924) 355; Burkill, Dict. Econ. Prod. Malay Penins.
(1935) 251.
Tree up to 40 m tall, sometimes with buttresses, evergreen. Leafy twigs 1.5 – 4 mm

thick, (sparsely) whitish to brownish appressedly puberulous, smooth, drying pale to
dark brown. Leaves distichous; lamina (sub)coriaceous, entire or pinnatifid to 3 – 5-
lobate when juvenile, elliptic to (sub)ovate, (2 –)10 – 20(– 33) by (1–)4 –12(–16) cm,
apex acuminate, base cuneate to rounded (occasionally peltate with a narrow strip of
laminar tissue connecting both sides), margin entire, ± revolute; upper surface (sub)-
glabrous, smooth; lower surface densely minutely whitish puberulous to tomentose
in the areoles or also on the reticulum, to sparsely whitish appressedly puberulous to
subtomentose on the main veins, smooth, glaucous when fresh; lateral veins (6 –)8 –15
pairs, rarely forked away from the margin, tertiary venation scalariform, ± prominent;
petiole (0.5 –)1– 2.5(– 4.5) cm long, 1– 2.5 mm thick, sparsely to densely whitish appres
sedly puberulous, the epidermis persistent; stipules lateral, 0.3–0.5 cm long, brownish
appressedly puberulous, caducous. Staminate inflorescences axillary, solitary or in pairs;
peduncle 1– 3 mm long, minutely brownish puberulous; head cylindrical to clavate to
ellipsoid, (5 –)10 –15(– 25) mm long, 2 – 4 mm diam.; perianth 2- or 3-parted, c. 0.5
mm long; stamen c. 0.8 mm long, anther c. 0.2 mm long; interfloral bracts (sub)peltate,
the apical part c. 0.4 mm diam., puberulous. Pistillate inflorescences axillary, solitary;
peduncle 2 – 5 mm long, densely brown puberulous; head subglobose; stigma simple;
interfloral bracts peltate, the apical part 0.2 – 0.3 mm diam., puberulous. Infructescences
subglobose, up to 7 cm diam., densely whitish to brownish puberulous (to velutinous);
Distribution — Australia (Northern Territory); in Malesia: Sumatra, Malay Penin-
sula, Borneo, Java, Lesser Sunda Islands (Flores, Sumba, Roma), Moluccas (Tanimbar
Islands).
Notes — 1. This species is characterised by the minute indumentum in the areoles.
2. The base of the lamina is occasionally ‘peltate’ (see p. 108): Ambri & Arifin W882
(L) from East Kalimantan.
5. Artocarpus gomezianus Wall. ex Trécul

Artocarpus gomezianus Wall. ex Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 118; King in Hook.f., Fl.
Brit. India 5 (1888) 544; Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 15, t. 14A; Burkill, Dict. Econ.
Prod. Malay Penins. (1935) 253; Kanjilal et al., Fl. Assam 4 (1940) 269; F.M. Jarrett, J. Arnold
Arbor. 41 (1960) 88; Backer & Bakh.f., Fl. Java 2 (1965) 19; Mabb., Taxon 26 (1977) 535; Kochum-
men, Tree Fl. Malaya 3 (1978) 126, t. 4; Corner, Wayside Trees Malaya ed.3, 2 (1988) 517.
Artocarpus petiolaris Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 422.
Artocarpus pomiformis Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indië 25 (1863) 400; Koord. &
Valeton, Bijdr. Boomsoort. Java 11 (1906) 23; J.J. Sm., Icon. Bogor. 3 (1907) 87, t. 235; Koord.,
Exkurs.-Fl. Java 2 (1912) 94.
Tree up to 40 m tall, evergreen. Leafy twigs 2 – 4 mm thick, sparsely whitish appress-

edly puberulous, smooth, drying pale to dark brown. Leaves distichous; lamina coria-
ceous, entire, elliptic, 10 – 25 by 6 –16 cm, apex acuminate (mostly shortly and abruptly
so) to rounded, base cuneate (to subattenuate) to rounded to subcordate, slightly unequal
to equal, margin entire, ± revolute; upper surface sparsely minutely whitish appress-
edly puberulous on the midrib or subglabrous, smooth; lower surface sparsely minutely
whitish appressedly puberulous on the midrib or main veins, smooth (to scabridulous);
lateral veins 7–10(–15) or 10 –15(– 20) pairs, mostly some of them (in the upper part
of the lamina) forked away from the margin, tertiary venation scalariform, slightly
prominent; petiole (1–)1.5 – 3 cm long, 1.5 – 2.5 mm thick, sparsely minutely whitish ap-
pressedly puberulous, the epidermis persistent; stipules lateral, 0.3 – 0.5 cm long, white
appressedly puberulous, caducous. Staminate inflorescences axillary, solitary; peduncle
5 –15 mm long, minutely white puberulous; head obovoid to subglobose, 10 –15 mm
diam.; perianth 2- or 3-parted, c. 0.5 mm long; stamen c. 0.6 mm long, anther c. 0.2
mm long; interfloral bracts (sub)peltate, the apical part c. 0.3 mm diam., puberulous.
Pistillate inflorescences axillary, solitary; peduncle 15 – 45 mm long, densely minutely
whitish to brownish puberulous; head obovoid to subglobose; stigma simple; interfloral
bracts peltate, caducous, the apical part c. 0.3 mm diam., puberulous. Infructescences
subglobose, up to 8 cm diam., densely whitish to brownish puberulous (to velutinous);
fruits ellipsoid, c. 1.2 cm long.
Distribution — India (Andaman Islands), Myanmar, Thailand, Indochina; in Malesia:
Sumatra, Malay Peninsula, Java, Philippines (Luzon, Cagayan Sulu).
Habitat — Evergreen and semi-evergreen forest at low altitudes.
Notes — 1. In Sri Lanka and India the species is also represented by a form with
dense whitish indumentum on the lamina, petiole and leafy twig (subsp. zeylanicus F.M.
Jarrett, J. Arnold Arbor. 41 (1960) 90).
2. As the species is not known from Borneo, Jarrett (1941) suggested that it was
introduced in Cagayan Sulu, but a recent collection (Ridsdale et al. 1394) from Luzon
may indicate that the species is indigenous. The fact that the leaves of material from the
Philippines normally have fewer lateral veins (7–10 pairs, whereas elsewhere normally
10 –15 pairs) would support the latter view.
6. Artocarpus lacucha Buch.-Ham.

Artocarpus lacucha Buch.-Ham., Mem. Wern. Nat. Hist. Soc. 5 (1826) 333; Mabb., Taxon 26 (1977)
529; A.J.C. Grierson & D.G. Long, Fl. Bhutan 1, 1 (1983) 100. — Artocarpus lakoocha Roxb., Fl.
Ind. 3 (1832) 524; [Hort. Bengal. (1814) 66 nomen, ‘lacucha’]; Graham, Cat. Pl. Bombay (1839)
193; Wight, Icon. Pl. Ind. Orient. 2 (1843) t. 681; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 117;
Kurz, Forest Fl. Burma 2 (1877) 433; King in Hook.f., Fl. Brit. Ind. 5 (1888) 543; Ann. Roy. Bot.
Gard. (Calcutta) 2 (1889) 14, t. 13; Renner, Bot. Jahrb. Syst. 39 (1907) 370; Burkill, Dict. Econ.
Prod. Malay Penins. (1935) 257.
Artocarpus ovatus Blanco, Fl. Filip. (1837) 666; ed. 3, 3 (1879) 73, plate; S. Vidal, Revis. Pl. Vasc.
Filip. (1886) 254; Ahern, Timber Tree Sp. Philip. (1901) 21, plate; F.M. Jarrett, J. Arnold Arbor.
41 (1960) 84.
Artocarpus cumingiana Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 119, t. 117, 118; Fern.-Vill., Nov.
App. (1880) 203; Elmer, Leafl. Philipp. Bot. 2 (1909) 620; Whitford, Bull. Bur. Forest. Philipp.
Islands 10, 2 (1911) 28, t. 6; Merr., Sp. Blancoan. (1918) 125; W.H. Br., Min. Prod. Philip. For. 2
(1921) 70, f. 23; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 40.
Artocarpus dadah Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 420; Ann. Mus. Bot. Lugduno-Batavi 3
(1867) 213; S. Moore, J. Bot. 63, Suppl. (1925) 112; Corner, Gard. Bull. Singapore 10 (1939) 282;
Wayside Trees Malaya (1940) 653; F.G. Browne, For. Trees Sar. & Brunei (1955) 352; F.M. Jarrett,
J. Arnold Arbor. 41 (1960) 91; Kochummen, Tree Fl. Malaya 3 (1978) 126, t. 4; Corner, Wayside
Trees Malaya ed. 3, 2 (1988) 516; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 194.
Artocarpus mollis Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 420.
Artocarpus rufescens Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 420.
Artocarpus tampang Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 421.
Ficus tampang Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 425.
Ficus inconstantissima Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 431. — Artocarpus inconstantissima
(Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 211.
Artocarpus fretessii Teijsm. & Binn. in Hassk., Abh. Naturf. Ges. Halle 9 (1866) 189, ‘fretissi’; Merr.,
Interpr. Herb. Amboin. (1917) 191; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 103. — Antiaris fretessii
Teijsm. & Binn., Cat. Hort. Bot. Bogor. (1866) 84, nomen.
Artocarpus dadah Miq. var. pubescens Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 213.
Artocarpus lakoocha Roxb. var. malayana King in Hook.f., Fl. Brit. India 5 (1888) 543; Ann. Roy.
Bot. Gard. (Calcutta) 2 (1889) 15; Ridl., Fl. Malay Penins. 3 (1924) 355; F.M. Jarrett, J. Arnold
Arbor. 41 (1960) 113.
Artocarpus refracta Becc., For. Borneo (1902) 630. — Artocarpus vrieseanus Miq. var. refractus
(Becc.) F.M. Jarrett, J. Arnold Arbor. 41 (1960) 98.
Artocarpus reniformis Becc., For. Borneo (1902) 631.
Prainea rumphiana Becc., For. Borneo (1902) 636. — [Metrosideros spuria Rumph., Herb. Amboin.
3 (1743) 26, t. 13.]
Artocarpus dasyphylla Miq. var. flava J.J. Sm., Icon. Bogor. 3 (1907) 85, t. 234.
Artocarpus leytensis Elmer, Leafl. Philipp. Bot. 1 (1908) 279; 2 (1909) 622; Merr., Enum. Philipp.
Flow. Pl. 2 (1923) 42.
Artocarpus paloensis Elmer, Leafl. Philipp. Bot. 1 (1908) 280; 2 (1909) 621.
Artocarpus acuminatissima Merr., Philipp. J. Sci. 18 (1921) 49; Enum. Philipp. Flow. Pl. 2 (1923) 40.
Artocarpus peltatus Merr., J. Straits Branch Roy. Asia. Soc. 85 (1922) 166; F.M. Jarrett, J. Arnold
Arbor. 41 (1960) 92; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 205, t. 4.
Artocarpus cumingiana Trécul var. stenophylla Diels, Bot. Jahrb. Syst. 67 (1935) 177.
Artocarpus yunnanensis Hu, Bull. Fan Mem. Inst. Biol. Bot. 8 (1937) 32.
Artocarpus ficifolia W.T. Wang, Acta Phytotax. Sin. 6 (1957) 274, t. 15, f. 23.
Artocarpus vrieseanus Miq. var. papillosus F.M. Jarrett, J. Arnold Arbor. 41 (1960) 99.
Artocarpus dasyphylla auct. non Miq.: Merr., Pl. Elmer. Born. (1929) 46.
Tree up to 40 m tall, deciduous. Leafy twigs 1.5 – 5(– 8) mm thick, densely brown to
yellowish (patent to appressedly) puberulous often intermixed with longer brown unci-
nate hairs, smooth, drying brown (to blackish); with or without up to 5 mm long short-
shoots on the older wood. Leaves distichous; lamina subcoriaceous to chartaceous,
entire or pinnatifid to 3-lobate when juvenile, basally attached (or occasionally peltate),
elliptic (to subrotundate), (3 –)10 – 30(– 33) by (2 –)6 –17 cm, apex acuminate, base cor-
date to subcordate to rounded (to obtuse to subcuneate), often slightly unequal, margin
denticulate towards the apex, particularly the acumen, sometimes entire, ± revolute (at
least towards the base); upper surface brownish to whitish puberulous to subtomentose
(mainly) on the main veins, smooth (occasionally ± scabrous or ± minutely bullate);
lower surface brownish to whitish patent to appressedly puberulous to subtomentose
or to hirtellous on the veins, smooth (to scabridulous); lateral veins (6 –)10 – 20 pairs,
in the lower part of the lamina usually faintly loop-connected, the lower ones spaced
or ± crowded (2 or 3 pairs departing close together), none (or sometimes one or a few)
branched or forked away from the margin, tertiary venation scalariform, ± prominent;
petiole (0.5 –)0.8 – 2(– 3) cm long, 1.5 – 3(– 4) mm thick, brown puberulous and often
intermixed with longer uncinate hairs, the epidermis persistent (or flaking off); stipules
lateral, 0.3 – 0.5(–1) cm long, brownish to whitish puberulous, caducous (or subpersist-
ent), or persistent, at the shoot-apices forming the terminal buds (or also in the leaf
axils and forming the lateral buds). Staminate inflorescences axillary (or just below the
leaves), solitary, or 1– 5 on short-shoots on the older wood, patent or deflexed; peduncle
3 – 35 mm long, densely brown puberulous; head subglobose to ellipsoid, to clavate (or
to pulvinate), to 3 cm long, 3 –15 mm diam.; perianth with 2 or 3 free tepals; stamen
0.5 – 0.8 mm long, anther 0.1– 0.2 mm long; interfloral bracts peltate, the apical part
c. 0.3 mm diam., puberulous, caducous or persistent. Pistillate inflorescences axillary
or below the leaves (down to previous season’s growth), solitary; peduncle (5 –)10 – 80
mm long, minutely brown velutinous; head subglobose to obovoid; perianth minutely
puberulous; stigma simple; interfloral bracts peltate, caducous (or persistent), the api-
cal part c. 0.3 mm diameter. Infructescences to c. 7 cm diam., often ± lobed, green to
orange with pink to red flesh, surface minutely velutinous; fruits ellipsoid, 1–1.2 cm
long. — Fig. 20a–e.
Distribution — Bhutan, India, Bangladesh, China (Yunnan), Myanmar, Thailand,
Indochina, Solomon Islands; in Malesia: Sumatra, Malay Peninsula, Borneo, Philip-
pines, Celebes, Moluccas (incl. Aru Islands), New Guinea.
Habitat — Mostly in evergreen forest at low altitudes. In New Guinea up to 2000 m.
Notes — 1. The species as currently defined includes material that Jarrett (J. Arnold
Arbor. 41 (1960) 73 –136) placed in A. dadah, A. fretessii, A. ovatus, A. vrieseanus
var. papillosus and var. refractus, as well as A. peltatus, which was separated from
A. dadah in the Flora Malesiana manuscript. The species can be recognised by features
indicating intermittent growth (in combination with deciduousness) such as the pres-
ence of terminal buds formed by some persistent stipules, the scars of which in addi-
tion to colour differences mark previous seasons’ growth of branches from the current
growth. The stipules are often subpersistent on opening-shoots and relatively long, to
1 cm. The lamina is chartaceous to subcoriaceous and the margin is denticulate at the
apex, sometimes only at the acumen. The base of the lamina is more or less unequal,
and varies from cordate through subcordate, rounded, and obtuse to cuneate. The latter
state occurs most frequently in the northern part of the species range. The lamina varies
in the lateral veins: the lower veins either distant or 2 or 3 pairs departing the midrib
close together as often seen in the eastern part of the species range. Patterns of variation
in length, direction, and colour of the indumentum are less clear, but the hairs are often
more or less appressed, shorter and paler in the eastern part of the species range. The
areoles of the lower surface are sometimes minutely bullate in the Malay Peninsula,
more frequently further northwards. In Borneo the base of the lamina is sometimes
peltate (with a narrow strip of lamina tissue connecting the two sides of the base of the
lamina). This was the distinguishing character of A. peltatus.
2. The pistillate inflorescences vary in length of the peduncle, the shedding of the
interfloral bracts, and the surface of the head in fruiting state which ranges from smooth
to finely ribbed or sometimes with the free parts of the apices of the pistillate flowers
short-pyramidate and hardened. Long peduncles, up to 8 cm long are found in northern
9 cm
3.5 cm
6 cm a
h
9 cm
g 2.5 cm
f
b
6 mm
9 cm d e c 1 cm
Fig. 20. a – e: Artocarpus lacucha Buch.-Ham. (‘fretissii’-form). a. Leafy twig; b. leafy twig with pistil-
late inflorescences; c. pistillate inflorescences; d. leafy twig with staminate inflorescences; e. staminate
inflorescence. — f– h: Artocarpus reticulatus Miq. f. Pistillate inflorescence; g. pistillate inflorescence;
h. staminate inflorescence (a: Kostermans s.n. Hort. Bog.; b, c: Cel. I-19; d, e: Cel. III-55; f: De Vriese
s.n., Celebes; g, h: Cel. V-228).
Borneo (in material that was identified as A. peltatus) and in the Philippines (in mate-
rial identified as A. ovatus). The peduncle becomes mostly up to 4 or 4.5 cm long. It
is often shorter in the Asian mainland (in material that was identified as A. lacucha by
Jarrett 1960). The interfloral bracts are often absent in the fruiting state and may be
already shed at anthesis. However, infructescences with persistent apices can be found
throughout the range of the species.
3. The staminate inflorescences vary more in their position, direction, length of the
peduncle, and shape and size of the head. Long peduncles, up to 4.5 cm long, are cor-
related with long peduncles of pistillate inflorescences. They are often deflexed in the
Philippines. The inflorescences are normally axillary or just below the leaves, mostly
solitary, sometimes in pairs, but often (also) borne on short-shoots on the older wood in
Celebes and the Moluccas, less often in the Philippines and (western) New Guinea. The
head varies considerably in shape, from subglobose to ellipsoid to clavate, or sometimes
to pulvinate or spicate, and ranges in size from c. 0.3 cm in length or diameter, to 4 cm
long in clavate to spicate ones or to 1.5 cm in diameter in subglobose ones.
4. The following informal entities can be distinguished:
‘lacucha’-form with the apex of the lamina often abruptly short-acuminate, the base of
the lamina often subcuneate and clearly unequal. Staminate inflorescences axillary,
solitary; peduncle 2 – 5 mm long; head ellipsoid to obovoid (to clavate) 1– 2.5 cm
long, 0.5 –1.5 cm diameter. Pistillate inflorescences: peduncle to 1.5(– 2.5) cm long;
interfloral bracts caducous or persistent, the free apices of the pistillate flowers
sometimes short-pyramidate and hardened. — India, (north-eastern & Andaman
Islands), Bangladesh, Myanmar, Thailand, China (Yunnan), Indochina. — In ever-
green, semi-evergreen or moist deciduous forest mostly at low altitudes.
‘dadah’-form with the base of the lamina mostly obtuse to subcordate, slightly un-
equal. Staminate inflorescences axillary (or just below the leaves) solitary, patent;
peduncle 0.4 –1.5 cm long; head subglobose (to pulvinate), 0.4 –1.5 cm diameter.
Pistillate inflorescences: peduncle to 3.5 cm long; interfloral bracts usually (early)
caducous; free parts of apices of pistillate flowers occasionally short-pyramidate
and hardened; surface of syncarp usually finely ribbed. — Myanmar (?), Thailand,
Sumatra, Malay Peninsula, Borneo. — In forest at low altitudes.
‘peltatus’-form with the base of the lamina rounded to subcordate, slightly unequal,
sometimes peltate. Staminate inflorescences axillary or just below the leaves, soli-
tary, patent; peduncle 2.5 – 3.5 cm long; head ellipsoid to subglobose, 0.7– 2 cm
diameter. Pistillate inflorescences: peduncle up to 8 cm long; heads of interfloral
bracts caducous; surface of ‘syncarp’ usually finely ribbed. — Borneo (northern).
— In evergreen forest at altitudes up to 1000 m.
‘ovatus’-form with the base of the lamina usually cordate to subcordate, slightly un-
equal. Staminate inflorescences axillary, below the leaves, solitary, paired, or some-
times up to 5 together on short-shoots on the older wood, often deflexed; peduncle
(0.5 –)1– 4.5 cm long; head obovoid to clavate, 1– 3 cm long, 0.3 –1.3 cm diam.;
interfloral bracts persistent. Pistillate inflorescences: peduncle up to 8 cm long,
deflexed at anthesis (?); interfloral bracts persistent. — Philippines. — In forest
at low altitudes.
The ‘peltatus’-form and ‘ovatus’-form are largely similar and distinguished by the
patent versus deflexed staminate inflorescences.
‘fretesii’-form with the base of the lamina usually rounded to cordate, slightly unequal.
Staminate inflorescences predominantly on short-shoots on the older wood, 1– 5
together, patent (or deflexed?); peduncle 0.3 – 0.7 cm long; head ellipsoid to subglo-
bose, 0.3 – 0.7 mm diameter. Pistillate inflorescences: peduncle to 4.5 cm long; head
subglobose to obovoid; interfloral bracts early caducous to persistent. — Celebes,
Moluccas. – At low altitudes.
‘refractus’-form with the base of the lamina usually rounded to cordate, slightly un-
equal. Staminate inflorescences usually axillary or just below the leaves, solitary,
or sometimes 1– 4 on short-shoots on the older wood, patent; peduncle 0.3 –1.2 cm
long; head subglobose 0.3 – 0.6 cm diam.; interfloral bracts persistent. Pistillate in-
florescences: peduncle to 5 cm long; interfloral bracts usually caducous; surface of
‘syncarp’ smooth (= not finely ribbed). — New Guinea, Solomon Islands. — Forest
at altitudes up to 2000 m.
The reticulum of the lamina lower surface is prominent in the few collections between
c. 1000 and 2000 m in New Guinea. This form also includes the type of A. vrieseanus
Miq. var. papillosus F.M. Jarrett. The epidermis of the petiole in this form sometimes
flakes off, whereas rarely in the forms treated above.
The ‘fretessii’-form and ‘refractus’-form are similar in the small subglobose, shortly
pedunculate staminate inflorescences. They differ in the frequency of staminate inflores-
cences on short-shoots on the older wood. The ‘refractus’-form grades into a ‘stenophyl-
lus’-form, comprising several collections which Jarret in her Flora Malesiana manuscript
called the eastern variants ‘stenophyllus α and β of A. vrieseanus and related to the
‘refractus’ variant’. In this form, the indumentum is relatively sparse and on leafy twigs
and lamina predominantly whitish and mostly intermixed with (± retrorse) uncinate hairs.
The petiole is usually 0.5 – 0.8 cm long and its epidermis exfoliates. The leafy twigs
are rather slender, mostly 2 – 3 mm thick, the trees are often less than 10 m tall without
buttresses, and no clear indication of intermittent growth. The peduncle of the pistillate
inflorescence is up to 5 cm long. — New Guinea, most frequently in the eastern part, but
also found in Biak, Yapen, and in the Solomon Islands. The sympatric occurrence of the
‘refractus’-form and the ‘stenophyllus’-form together with the nature of the differentiat-
ing characters cast doubt on the distinctiveness of the latter. Future studies with more
material are needed to clarify their taxonomic status. The ‘stenophyllus’-form resembles
A. nitidus from New Guinea and the Solomon Islands, but differs in the patent hairs on
the lamina lower surface and the usually denticulate margin towards the apex.
7. Artocarpus longifolius Becc.

Artocarpus longifolius Becc., For. Borneo (1902) 629; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 83;
Tree up to 20 m tall, of intermittent growth, with terminal (and lateral?) buds, de-
ciduous (?). Leafy twigs (2 –)4 – 6 mm thick, densely brownish minutely puberulous
and longer uncinate hairs, smooth, drying dark brown. Leaves distichous; lamina (sub)-
coriaceous, entire, elliptic, 9 – 33 by 5 –14.5 cm, apex (sub)acuminate, base obtuse to
rounded, margin entire, ± revolute; upper surface brownish puberulous on the main
veins or only on the midrib, smooth; lower surface ± densely to sparsely brownish to
whitish puberulous to hirtellous on the veins or to minutely puberulous on the smaller
veins, smooth (to scabridulous); midrib, lateral and tertiary veins mostly ± impressed,
lateral veins (5 –)8 –15 pairs, not forked away from the margin, tertiary venation scalari
form, prominent, the smaller veins also prominent; petiole 0.3 – 0.8 mm long, 3 – 4 mm
thick, brownish puberulous, the epidermis flaking off; stipules lateral, 0.3 – 0.5(–1) cm
long, brownish puberulous, caducous (or subpersistent). Staminate inflorescences axil-
lary, solitary; peduncle 1– 3 cm long, minutely brown puberulous and with numerous
longer brownish uncinate hairs; head subglobose, c. 6 mm diam.; perianth with 3 free
tepals, c. 0.4 mm long; stamen c. 0.8 mm long, anther 0.1– 0.2 mm long; interfloral
bracts peltate, the apical part c. 0.3 mm diam., puberulous. Pistillate inflorescences
axillary, solitary; peduncle 2 – 6 cm long, minutely brown puberulous and with numer-
ous longer brownish uncinate hairs; head subglobose to ellipsoid; perianth puberulous;
stigma simple; interfloral bracts peltate, persistent, the apical part 0.2 – 0.3 mm diameter.
Infructescences (immature) c. 2 cm diam., densely brown puberulous (velutinous); fruit
not seen.
Notes — 1. The species can be recognised by the short and thick petiole with exfo-
liating epidermis.
2. Jarrett (1960) assumed that subglabrous specimens represented (sub)juvenile
material. However, two such specimens are fertile and a specimen bearing a pistillate
inflorescence has a much shorter peduncle (c. 0.3 cm long as opposed to 2 – 6 cm long
in ± densely hairy specimens). The differences in indumentum possibly correlate with
the length of the peduncles of the pistillate inflorescences and suggest distinction of
subspecies, or closely related species. More material is needed to sort out the status of
these two morphological entities which are sympatric and apparently have a similar
ecology.
a. subsp. longifolius
Lower surface of the lamina ± densely brownish puberulous to hirtellous, partly with
uncinate hairs; midrib, lateral and tertiary veins mostly ± impressed above.
Habitat — Evergreen forest at low altitudes
b. subsp. adpressus C. C. Berg

Artocarpus longifolius Becc. subsp. adpressus C. C. Berg, Blumea 50 (2005) 543.
Lower surface of the lamina sparsely brownish to whitish appressed puberulous on the
main veins, this indumentum intermixed with patent to retrorse uncinate hairs, minutely
puberulous on the smaller veins; the main veins not (never?) impressed above.
Distribution — Borneo (Brunei, Central Kalimantan).
Note — In the collection from Brunei (Tutong, Telisai, Bukit Basong, N. Nangkat
et al. BRUN 15511), the leaves are small (9 –13 by 3.5 – 6 cm) and have few lateral
veins (5 –7 pairs), whereas in the collection from Central Kalimantan (Taman Wisata,
Bukit Tengkiling area, Palangka Raya, Abriansyah & Z. Arifin AA 954) the size of the
lamina varies from small (as in the Brunei collection) to the normal size for subsp.
longifolius.
8. Artocarpus nitidus Trécul

Artocarpus nitidus Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 119; Miq., Fl. Ned. Ind. 1, 2 (1859) 288;
Fern.-Vill., Nov. App. (1880) 203; S. Vidal, Revis. Pl. Vasc. Filip. (1886) 254; Elmer, Leafl. Philipp.
Bot. 2 (1909) 624; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 121; Corner, Wayside Trees Malaya ed.3,
2 (1988) 520, t. 157, 162; Kochummen, Tree Fl. Sabah & Sarawak 3 (2000) 203.
Artocarpus lanceolata Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 121.
Artocarpus gomezianus Wall. ex Trécul var. griffithii King in Hook.f., Fl. Brit. India 5 (1888) 544; Ann.
Roy. Bot. Gard. (Calcutta) 2 (1889) 15, t. 14B. — Artocarpus griffithii (King) Merr., Pap. Michi-
gan Acad. Sci. I, 24 (1939) 64. — Artocarpus nitidus Trécul subsp. griffithii (King) F.M. Jarrett,
J. Arnold Arbor. 41 (1960) 128; Kochummen, Tree Fl. Malaya 3 (1978) 131, t. 6.
Artocarpus humilis Becc., For. Borneo (1902) 629. — Artocarpus nitidus Trécul subsp. humilis (Becc.)
F.M. Jarrett, J. Arnold Arbor. 41 (1960) 126.
Artocarpus antiarifolia Becc., For. Borneo (1902) 630.
Artocarpus xanthocarpus Merr., Philipp. J. Sci. 17 (1904) 10; Philipp. J. Sci., 1, Suppl. (1906) 43;
Elmer, Leafl. Philipp. Bot. 2 (1909) 626; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 102.
Artocarpus borneensis Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 165. — Artocarpus nitidus
Trécul subsp. borneensis (Merr.) F.M. Jarrett, J. Arnold Arbor. 41 (1960) 127.
Artocarpus eberhardtii Gagnep., Bull. Soc. Bot. France 73 (1926) 87; Fl. Indo-Chine 5 (1928) 737.
Artocarpus parva Gagnep., Bull. Soc. Bot. France 73 (1926) 89; Fl. Indo-Chine 5 (1928) 735, t. 90.
Artocarpus sampor Gagnep., Bull. Soc. Bot. France 73 (1926) 90; Fl. Indo-Chine 5 (1928) 738.
Artocarpus eberhardtii Gagnep. var. poilanei Gagnep., Fl. Indo-Chine 5 (1928) 737.
Artocarpus lingnanensis Merr., Lingnan. Sci. J. 7 (1931) 302; 13 (1934) 56. — Artocarpus nitidus
Trécul subsp. lingnanensis (Merr.) F.M. Jarrett, J. Arnold Arbor. 41 (1960) 124.
Artocarpus vrieseanus Miq. var. subsessilis F.M. Jarrett, J. Arnold Arbor. 41 (1960) 101.
Artocarpus lamellosa auct. non Blanco: Merr., Philipp. J. Sci. 27 (1905) 80; Sp. Blancoan. (1918)
124; Enum. Philipp. Flow. Pl. 2 (1923) 41.
Artocarpus rubrovenius auct. non Warb.: Merr., Philipp. J. Sci., Bot. 3 (1908) 401.
Artocarpus lanceolata auct. non Trécul: Merr., Enum. Philipp. Flow. Pl. 2 (1923) 42; Gagnep. Fl.
Indo-Chine 5 (1928) 738.
Artocarpus gomezianus auct. non Wall. ex Trécul: Ridl., Fl. Malay Penins. 3 (1924) 355; Corner, Way
side Trees Malaya (1940) 654, t. 195.
Tree up to 35 m tall, sometimes with buttresses, evergreen. Leafy twigs (1–)2 – 3(– 4)
mm thick, (sparsely) whitish to brownish appressedly puberulous (sometimes also ret-
rorse uncinate hairs), smooth, drying brown (to blackish). Leaves distichous; lamina
(sub)coriaceous, entire, elliptic, (3 –)6 –18(– 33) by (1.5 –)3.5 – 9 cm, apex acuminate
with the acumen long or short, caudate, acute, obtuse (or to rounded), base cuneate to
subattenuate to rounded or to subcordate, equal to ± unequal, margin entire (or crenate
towards the apex, or when juvenile pinnately lobate), ± revolute (to flat); upper sur-
face glabrous or whitish (largely) appressedly puberulous on the main veins or only
on the midrib, often partly with retrorse uncinate hairs, smooth; lower surface sparsely
whitish to brownish puberulous on the (main) veins (the hairs mostly appressed or
some patent or retrorse and then often uncinate) to glabrous, smooth (or scabridulous);
lateral veins (5 –)7–10(–15) pairs, none of them forked away from the margin, tertiary
venation (sub)scalariform, almost flat to slightly prominent; petiole (0.5 –)8 –1.5(– 2.5)
cm long, 1–2 mm thick, sparsely whitish to brownish appressedly puberulous (some-
times also retrorse uncinate hairs), the epidermis persistent; stipules lateral, 0.2 – 0.4 cm
long, brownish to whitish appressedly puberulous, caducous. Staminate inflorescences
axillary, solitary; peduncle1– 4 mm long, densely brown to whitish puberulous; head
subglobose to obovoid to narrowly ellipsoid to clavate, 2 –12 by 2 – 5 mm; perianth 2-

or 3-lobed to -parted, c. 0.6 mm long; stamen c. 0.8 mm long, anther c. 0.2 mm long;
interfloral bracts (sub)peltate, the apical part c. 0.4 mm diam., puberulous. Pistillate
inflorescences axillary, solitary; peduncle 1– 5(– 25) mm long, densely brown puberu-
lous; head subglobose to ellipsoid; stigma simple; interfloral bracts peltate, persistent
or caducous, the apical part c. 0.3 mm diam., puberulous. Infructescences subglobose,
up to c. 3(– 4?) cm diam., subglabrous, sparsely hairy or densely hairy (velutinous);
Distribution — India (north-eastern), Myanmar, Thailand, China (southern, accord-
ing to Zhekun & Gilbert, Fl. China 5 (2003) 34 also in Lan Yu (Taiwan)), Indochina,
Solomon Islands; in Malesia: Sumatra, Malay Peninsula, Borneo, Philippines, Moluc-
cas (Aru Islands), New Guinea.
Habitat — Evergreen forest at altitudes up to c. 1600 m, in New Guinea up to 1750 m.
Notes — 1. The subspecies is rather variable and allows recognition of three informal
entities:
‘nitidus’-form – Base of the lamina tending towards subcordate, lateral veins mostly
5 or 6 pairs, epidermis of the petiole persistent; heads of the staminate inflores-
cences mostly elongate, surface of the infructescences densely hairy (velutinous),
interfloral bracts of the pistillate inflorescences sometimes early caducous. — Phil-
ippines.
‘griffithii’-form with the base of the lamina tending towards attenuate, lateral veins
mostly 7–10, sometimes up to 15 pairs, epidermis of the petiole persistent; heads
of the staminate inflorescences mostly globose, surface of the infructescences sub-
glabrous or (in Borneo) sometimes (rather) sparsely puberulous, interfloral bracts
of the pistillate inflorescences persistent. — Sumatra, Malay Peninsula, Borneo,
Philippines. One of the collections from Peninsular Malaysia (FRI 6941) is distinct
in its up to 1 cm long peduncle.
‘borneensis’-form with the base of the lamina and lateral veins and staminate inflo-
rescences as in the ‘griffithii’-form – but the indumentum on the leafy twigs and
petioles often denser and the epidermis of the petiole often flaking off; surface of
the infructescence densely brown to purplish puberulous (velutinous), interfloral
bracts of the pistillate inflorescence persistent. — Borneo (Brunei, Kalimantan
(eastern), Sabah, Sarawak). Two collections from eastern Kalimantan are distinct
in their peduncles which are up to 2.5 cm long.
‘subsessilis’-form with the base of the lamina cuneate to subcordate, in contrast to the
western forms the midrib of the lamina is ± densely hairy above and the venation
of the lamina often more prominent beneath; the peduncles are on average slightly
longer; those of the pistillate inflorescences can be up to 2 cm long and of the
staminate inflorescences up to 1.5 cm long. — Occurring disjunct from the main
range in New Guinea and the Solomon Islands. This form includes the types of
A. vrieseanus var. subsessilis and probably also of A. antiarifolia.
2. Some collections from the Philippines (Batan Island, Luzon) have caudate apices
of the lamina.
3. The staminate inflorescences are occasionally borne on short-shoots on the older

wood: see Van Royen 3591 (Vogelkop).
4. The material from the Solomon Islands differs somewhat from New Guinean
material in the hairs on the midrib above which are all straight and point towards the
apex of the lamina rather than partly uncinate and retrorse.
5. As the type of A. antiarifolia could not be (re)examined, it is not quite certain that
the name is correctly included in the synonymy.
6. The collection FRI 19234 indicated as Artocarpus ‘sp. B’ by Kochummen (Tree
Fl. Malaya 3 (1978) 135) is referred to this species with some doubt. It has all charac-
ters of A. nitidus (including the subglabrous head of the pistillate inflorescence of the
‘griffithii’-form) except for the dense brown indumentum on the leafy twig, petiole,
and the midrib of the lamina, in which it resembles A. lacucha. Collections with similar
indumentum have been made in northern and eastern Borneo (e.g., Endert 3191 and
Paie & Yeo S 38380) and can be referred to the ‘borneensis’-form.
9. Artocarpus reticulatus Miq.

Artocarpus reticulatus Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 213, ‘reticulata’; Renner, Bot.
Jahrb. Syst. 39 (1907) 369; Koord., Suppl. Fl. Celebes 2 (1922) t. 3; 3 (1922) 1; F.M. Jarrett, J.
Arnold Arbor. 41 (1960) 107.
Artocarpus dadah Miq. var. celebica Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 213.
Tree up to 30 m tall with short buttresses, evergreen. Leafy twigs 3 – 6 mm thick,

sparsely or towards the nodes more densely brownish minutely appressedly puberulous,
smooth, drying brown to greyish. Leaves distichous; lamina subcoriaceous, entire, el-
liptic to subobovate, 15 – 35 by 8 –16 cm, apex acuminate, base rounded to obtuse, broad
bases usually minutely auriculate, equal, margin entire, slightly revolute to flat; upper
surface sparsely brownish appressedly puberulous on the midrib, smooth, occasionally
± bullate; lower surface sparsely brownish appressedly puberulous on the midrib to
whitish hispidulous on the smaller veins, ± scabrous; lateral veins 10 –17 pairs, none (or
one) forked away from the margin, tertiary venation scalariform, ± prominent; petiole
1– 2.5 cm long, 2 – 3 mm thick, sparsely whitish puberulous, the epidermis persistent;
stipules lateral, 0.3 – 0.5 cm long, brownish appressedly puberulous, caducous. Stami
nate inflorescences axillary (or on axillary short-shoots), solitary or in pairs, patent;
peduncle 8 –13 mm long, densely brown appressedly puberulous; head obovoid to sub-
globose, c. 1.5 cm diam.; perianth with 3 or 4 free tepals, c. 0.2 mm long; stamen c. 0.5
mm long, anther c. 0.2 mm; interfloral bracts peltate, persistent, the apical part 0.4 – 0.6
mm diam., ciliolate. Pistillate inflorescences axillary, solitary; peduncle 2.5 – 4.5 cm
long, broadened at the apex, densely minutely brown puberulous; head subglobose to
ellipsoid; perianth brown puberulous; stigma simple, c. 1.5 mm long; interfloral bracts,
few, peltate, the apical part c. 0.3 mm diam., persistent. Infructescences to c. 6 cm diam.,
lobed, with pyramidate hardened apices of the perianths; fruit ellipsoid, c. 1.2 cm long.
— Fig. 20f – h.
Distribution — Celebes, Moluccas (Ternate, Ambon).
Notes — 1. This species can be recognised by the sparsely hairy laminas with ±
scabrous lower surface and often minutely auriculate base.
2. The species resembles some specimens of A. lacucha and many species of subg.
Artocarpus in the hardened pyramidate free apices of the perianth.
10. Artocarpus rubrovenius Warb.

Artocarpus rubrovenius Warb. in Perkins, Fragm. Fl. Philipp. 3 (1905) 166; Merr., Philipp. J. Sci., 1,
Suppl. (1906) 43; Elmer, Leafl. Philipp. Bot. 2 (1909) 622; Merr., Enum. Philipp. Flow. Pl. 2 (1923)
43; W.H. Br., Useful Pl. Philipp. (1941) 470; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 114.
Tree up to 15 m tall, evergreen (?). Leafy twigs 2 – 4 mm thick, sparsely brownish to

whitish appressedly puberulous, smooth, drying dark brown to blackish. Leaves dis-
tichous; lamina (sub)coriaceous, entire, elliptic, 10 – 26 by 5 –14 cm, apex acuminate,
base cuneate to rounded, usually unequal, margin entire (or crenate to subdenticulate),
usually flat; upper surface sparsely brownish to whitish appressedly puberulous on the
main veins, smooth; lower surface sparsely brownish to whitish appressedly puberulous
on the main veins, smooth; lateral veins (6 –)7–10(–11) pairs, none or sometimes one
forked away from the margin, tertiary venation (loosely) scalariform, slightly promi-
nent; petiole (1–)1.5 – 2.5(– 3) cm long, 1– 2 mm thick, sparsely brownish to whitish
appressedly puberulous, the epidermis persistent; stipules lateral, 0.3 – 0.5 cm long,
brownish to yellowish appressedly puberulous, caducous. Staminate inflorescences
axillary, solitary (or in pairs); peduncle 2 – 5 mm long, brown short-velutinous; head
obovoid to ellipsoid, 1.5 – 3(– 4.5) cm long, 1–1.5(– 2) cm diam.; perianth with 2 – 4
free or basally connate tepals, c. 0.5 mm long; stamen c. 0.7 mm long, anther c. 0.2
mm long; interfloral bracts peltate, the apical part c. 0.3 mm diam., puberulous. Pistil
late inflorescences axillary, solitary; peduncle 0.4 –1 cm long, brown short-velutinous;
head ellipsoid to subglobose; stigma simple; interfloral bracts peltate, persistent, the
apical part c. 0.3 mm diam., puberulous. Infructescences subglobose, up to 8 cm diam.,
puberulous; fruit not seen.
Distribution — Philippines.
Notes — 1. This species can be distinguished from the sympatric A. nitidus by the
larger lamina and flower heads.
2. The trees may show intermittent growth.
11. Artocarpus subrotundifolius Elmer

Artocarpus subrotundifolius Elmer, Leafl. Philipp. Bot. 2 (1909) 619; Merr., Enum. Philipp. Flow. Pl.
2 (1923) 43; F.M. Jarrett, J. Arnold Arbor. 41 (1960) 108.
Tree up to 25 m tall of intermittent growth with terminal and axillary buds, deci
duous (?). Leafy twigs 4 – 6 mm thick, densely brown puberulous to velutinous, smooth,
drying brown to blackish. Leaves distichous; lamina (sub)coriaceous, entire, elliptic
to narrowly elliptic or to subrotundate, 18 – 36 by 11– 27 cm, apex short-acuminate (to
rounded), base cordate to rounded, usually ± unequal to equal, margin entire to denticu-
late towards the apex, ± revolute to flat; upper surface brownish to whitish puberulous
mainly on the main veins, smooth; lower surface brownish to whitish puberulous to sub-
hispidulous on the veins, scabridulous; lateral veins 10 –14 pairs, most of them forked
or branched away from the margin, tertiary venation scalariform, prominent, also the
smaller veins prominent; petiole (1.5 –)3 –7 cm long, 2.5 – 3 mm thick, densely brown
puberulous to velutinous, the epidermis persistent; stipules lateral, 0.3 – 0.5 cm long,
brownish puberulous, caducous. Staminate inflorescences axillary, solitary (or in pairs);
peduncle (0.5 –)1.5 – 2.2 cm long, brown short-velutinous; head obovoid to ellipsoid to
subglobose, (2 –)2.5 – 5 cm long, 2 – 3.5 cm diam.; perianth with 3 or 4 free tepals, c. 0.6
mm long; stamen 1 mm long, anther c. 0.2 mm long; interfloral bracts peltate, the apical
part 0.3 – 0.5 mm diam., puberulous. Pistillate inflorescences axillary, solitary; peduncle
2 –7 (?) cm long, brown short-velutinous; head subglobose; stigma simple; interfloral
bracts peltate, the apical part 0.3 – 0.5 mm diam., puberulous, caducous. Infructescences
subglobose, up to 6 cm diam., puberulous; fruits subglobose, c. 1.5 diameter.
Distribution — Philippines.
Notes — 1. The species can be recognised by the large lamina with the lateral veins
frequently forked or branched away from the margin.
2. The species is probably closely related to A. tomentosulus.
12. Artocarpus tomentosulus F.M. Jarrett

Artocarpus tomentosulus F.M. Jarrett, J. Arnold Arbor. 41 (1960) 117; Kochummen, Tree Fl. Sabah
& Sarawak 3 (2000) 212.
Artocarpus primackii Kochummen, Gard. Bull. Singapore 50 (1998) 199; Tree Fl. Sabah & Sarawak
3 (2000) 207.
Tree up to 30 m tall of intermittent growth with terminal and axillary buds, deciduous
(?). Leafy twigs 2.5 – 4.5 mm thick, densely brownish to whitish minutely puberulous,
smooth, drying brown. Leaves distichous; lamina (sub)coriaceous, entire, elliptic, 10 – 30
by 5 – 20 cm, apex acuminate, base obtuse to rounded to subcord(ul)ate or to cuneate,
margin entire, ± revolute; upper surface whitish to brownish minutely puberulous on the
main veins, smooth; lower surface ± densely whitish to brownish minutely puberulous
to hispidulous on the veins, smooth to scabridulous; lateral veins (10 –)12 –16 pairs,
none forked away from the margin, tertiary venation scalariform, prominent; petiole
1– 4.5 cm long, 2 – 3 mm thick, densely whitish to brownish minutely puberulous, the
epidermis flaking off; stipules lateral, 0.3 – 0.5 cm long, brownish appressedly puberu-
lous, caducous. Staminate inflorescences axillary, solitary; peduncle c. 2 mm long,
brown velutinous; head subglobose, c. 6 mm diam.; perianth tubular, 2-lobed; stamen
not exserted, anther 0.1– 0.2 mm long; interfloral bracts peltate, the apical part c. 0.3
mm diam., puberulous. Pistillate inflorescences axillary, solitary on short-shoots with
buds lateral over the base of the peduncle; peduncle up to 4 cm long, brown velutinous;
head subglobose; perianth puberulous; stigma simple; interfloral bracts peltate, persist-
ent, the apical part 0.2 – 0.3 mm diam., persistent. Infructescences up to 6 cm diam.,
pinkish red at maturity.
Note — The species is characterised by the dense minute indumentum on the leafy
twig, petiole, lower surface of the lamina, and peduncle. The inflorescences, at least
the pistillate ones, are borne on short-shoots with distinct buds, consisting of several
persistent stipules.
13. Artocarpus vrieseanus Miq.

Artocarpus vrieseanus Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 212; F.M. Jarrett, J. Arnold
Arbor. 41 (1960) 95.
Tree up to 6 (or more?) m tall, evergreen. Leafy twigs 1.5 – 3 mm thick, sparsely whit-
ish appressedly puberulous to subglabrous, smooth, drying brown to blackish. Leaves
distichous; lamina subcoriaceous, entire, subobovate to elliptic, 7– 23 by 3.5 –11 cm,
apex acuminate, base rounded to obtuse (to cuneate), often ± unequal, margin entire,
slightly revolute to flat; upper surface (very) sparsely whitish appressedly puberulous
on the main veins, smooth; lower surface very sparsely whitish appressedly puberulous
to subglabrous on the main veins, smooth; lateral veins 9 or 10 pairs, none forked away
from the margin, tertiary venation scalariform, slightly prominent; petiole 1– 2.2 cm
long, 1– 2 mm thick, sparsely whitish appressedly puberulous, the epidermis persistent;
stipules lateral, 0.3 – 0.6 cm long, whitish to brownish appressedly puberulous, cadu-
cous. Staminate inflorescences axillary, just below the leaves (or on short-shoots on
the older wood), solitary (or in pairs), deflexed (?); peduncle 1.2 –1.7 cm long, densely
minutely whitish to brownish puberulous; head obovoid to subglobose, 3 – 5 mm diam.;
perianth 2- or 3-lobed, c. 0.5 mm long; stamen c. 0.8 mm long, anther 0.1– 0.2 mm
long; interfloral bracts peltate, the apical part 0.2 – 0.3 mm diam., sparsely puberulous.
Pistillate inflorescences axillary, solitary; peduncle 2.5 – 5.5 or 0.3 – 2 cm long, densely
minutely brownish or whitish puberulous; head globose; perianth puberulous; stigma
simple; interfloral bracts peltate, caducous or persistent, the apical part 0.2 – 0.3 mm
diam., puberulous. Infructescences to c. 4.5 cm diam., orange, subglabrous or sparsely
to densely minutely brownish to whitish puberulous (to subpapillate); fruit subglobose,
c. 1 cm diam.
Distribution — Celebes (Manado), Moluccas (Batjan).
Habitat — Evergreen forest, at altitudes up to 1000 m.
Note — Artocarpus vrieseanus as defined by Jarrett (1960) is too heterogeneous.
The material of var. papillosus and var. refractus is included in A. lacucha (see p. 115,
119) and that included in var. subsessilis is transferred to A. nitidus (see p. 121, 122).
Hullettia
Hullettia King ex Hook.f., Fl. Brit. India 5 (1888) 547; King, Ann. Roy. Bot. Gard. (Calcutta) 5, 2
(1896) 163, t. 197; Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 1 (1897) 122; Boerl., Handl.
Fl. Ned. Ind. 3 (1900) 338; Ridl., Fl. Malay Penins. 3 (1924) 358; F.M. Jarrett, J. Arnold Arbor.
41 (1960) 334.
Shrubs to small trees, monoecious. Leaves spirally arranged; lamina pinnately

veined, entire; stipules free, lateral, subulate, subpersistent. Inflorescences unisexual,
solitary or in pairs in the leaf axils, discoid- to turbinate-capitate, pedunculate, with
a row of bracts at the margin of the receptacle (interfloral bracts present in staminate
inflorescences, absent or rudimentary in pistillate ones?). Staminate flowers up to c. 30,
completely fused and perianths indistinct; stamens 2, filaments partly connate. Pistil
late flowers 2 – 6, completely fused and the perianths indistinct; ovaries free, stigma 1,
capitate. Infructescences subglobose, fleshy, orange to yellow; pericarp crustaceous;
seed without endosperm, embryo straight, longitudinally aligned, cotyledons equal,
radicle basal, short.
Distribution — The genus comprises two closely related species and ranges from
Myanmar to Sumatra.
Morphology — The discoid and involucrate inflorescences are reminiscent of the
type found in some members of the Castilleae and species of Dorstenia, in which genus
one of the species has been described. Small ovate to linear structures found among
staminate flowers of H. dumosa probably represent interfloral bracts. Minute elevations
with slightly different indumentum among the flowers of pistillate inflorescences may
also represent such bracts, but quite rudimentary.
1. Hullettia dumosa King ex Hook.f.

Hullettia dumosa King ex Hook.f., Fl. Brit. India 5 (1888) 547; King, Ann. Roy. Bot. Gard. (Calcutta)
5, 2 (1896) 163, t. 197; Ridl., Fl. Malay Penins. 3 (1924) 358; Burkill, Dict. Econ. Prod. Malay
Penins. (1935) 1202; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 9, f. 2g–k; 41 (1960) 337; Kochum-
men, Tree Fl. Malaya 3 (1978) 162, t. 8.
Shrub or tree up to 10 m tall. Leafy twigs (2 –)3 – 6 mm thick, densely puberulous.

Leaves spirally arranged; lamina (narrowly) obovate to elliptic (to narrowly elliptic),
5 mm
2 cm
b f
4 mm 1 cm
2 mm c 1 mm
d
h e
a
g
4 cm
Fig. 21. Hullettia dumosa King ex Hook.f. a. Leafy twig with pistillate inflorescences and infructes-
cence; b. pistillate inflorescence; c. pistillate inflorescence in length section; d. ovary in length sec-
tion; e. pistillate inflorescence in tangential section; f. infructescences in length section; g. staminate
inflorescences; h. part of staminate inflorescence in length section (a: Alvins 3290; b, d: Cameron s.n.
4-XI-1942; c, e: Ridley s.n. II–1921; f: Curtis 2404; g, h: Robinson s.n. 20-III-1913).
8 – 30(– 40) by 3 – 9(–12) cm, coriaceous, apex acuminate, base rounded to cuneate,
margin (sub)entire; upper surface glabrous; lower surface glabrous or very sparsely
minutely puberulous on the midrib, ± scabrous or smooth; midrib ± prominent above,
lateral veins 8 –12(–15) pairs, tertiary venation reticulate (to subscalariform); peti-
ole 1– 3.5(– 5) cm long, sparsely puberulous; stipules 0.3 –1(–1.5) cm long, subulate,
sparsely puberulous, subpersistent. Staminate inflorescences axillary, solitary or paired;
peduncle 1– 2 cm long, puberulous; receptacle discoid, c. 1 cm diam., puberulous out-
side, involucral bracts marginal, ovate, 3 – 4 mm long, puberulous; perianth c. 1.5 mm
long, puberulous; stamens c. 3 mm long, filaments connate, anthers c. 1 mm long;
interfloral bracts ovate to linear, puberulous. Pistillate inflorescences axillary, solitary
or paired; peduncle 1.5 – 4.5(– 5.5) cm long, puberulous; receptacle discoid, c. 1 cm
diam., puberulous outside; involucral bracts ovate, 2 – 3 mm long, puberulous; flowers
c. 8; perianth 1–1.5 mm long, apex densely minutely puberulous; stigmas clavate, 0.5 –1
mm long. Infructescences subglobose to discoid, up to 5.5 cm diam., with 1– 3 fruits,
densely puberulous to hispidulous, often ± scabrous, orange at maturity. — Fig. 21.
Distribution — Thailand; in Malesia: Sumatra, Malay Peninsula.
Habitat — Evergreen forest at altitudes up to c. 1000 m.
Note — Hullettia griffithii (Kurz) King, the other species, known from peninsular
Myanmar and Thailand, has not yet been recorded from Malesia. It is in many features
similar to H. dumosa, but it can easily be recognised by the midrib of the lamina being
slightly impressed above.
PARARTOCARPUS
Parartocarpus Baill., Adansonia 11 (1875) 294; Benth. & Hook.f., Gen. Pl. 3 (1880) 375; Engl. in
Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 82; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 335; Becc.,
For. Borneo (1902) 632; Renner, Bot. Jahrb. Syst. 39 (1907) 361; Becc., Webbia 5 (1923) 559; F.M.
Jarrett, J. Arnold Arbor. 41 (1960) 320; Corner, Gard. Bull. Singapore 19 (1962) 241; Backer &
Bakh.f., Fl. Java 2 (1965) 17; Corner, Gard. Bull. Singapore 28 (1976) 183, t. 1– 3.; Kochummen,
Tree Fl. Malaya 3 (1978) 162; Go, Tree Fl. Sabah & Sarawak 3 (2000) 318.
Gymnartocarpus Boerl., Icon. Bogor. 1 (1897) 73, t. 24, 25; Handl. Fl. Ned. Ind. 3 (1900) 335; Engl.
in Engl. & Prantl, Nat. Pflanzenfam. 2 (1900) 17; Koord. & Valeton, Bijdr. Boomsoort. Java 11
(1906) 28; Koord., Exkurs.-Fl. Java 2 (1912) 96; J.J. Sm., Bull. Jard. Bot. Buitenzorg III, 4 (1922)
233, t. 6 – 8.
Trees, dioecious. Leaves spirally arranged; lamina pinnately veined, entire; stip-
ules fused, semi-amplexicaul. Inflorescences unisexual (or rarely bisexual), solitary in
the leaf axils, pedunculate, with 3 – 8 large basally attached bracts at the base of the
head, interfloral bracts with free apices similar to the free parts of the tepals, dis-
coid to cushion shaped to aculeate. Staminate inflorescences (sub)globose or obovoid;
flowers numerous, connate; perianth tubular, free parts thickened, cushion shaped to
aculeate; stamens 1– 3, mostly 2, filaments free or partly connate. Pistillate inflores
cences (sub)globose; flowers numerous, connate except for the apices, apices discoid
to cushion shaped to aculeate; ovary free, stigma 1. Infructescences with the indurated
surface areolate or with cushion shaped to pyramidate to spinose processes; fruit with
a firm exocarp finally decaying and leaving the stony endocarp; seed with remains of
endosperm present, embryo curved, cotyledons incumbent and unequal, radicle long.
Distribution — The genus comprises two closely related species and ranges from
Thailand to the Solomon Islands.
Delimitation — This genus differs from Artocarpus in the interpetiolar connate stip-
ules, the well-developed basally attached bracts at the base of the head of the inflores-
cence, epigeal germination, and from most species of Artocarpus in the full connation
of perianths of the pistillate flowers.
Morphology — It is not clear whether all structures indicated as processes represent
the upper parts of tepals, or can be upper parts of interfloral bracts as well (see Jarrett,
J. Arnold Arbor. 40 (1959) 7). Those surrounding the style and the androecium, two to
four in number, are the free parts of perianths. Those not associated with style or sta-
mens may represent interfloral bracts. However, in the neotropical genus Naucleopsis
Miq., in which free parts of the tepals of pistillate flowers are aculeate to cushion shaped
and indurated, the processes that are not clearly associated with styles can be regarded
as derived from the perianth (see Berg, Fl. Neotrop. Monogr. 7 (1972) 107). In contrast
to Parartocarpus, there is a clear tendency of increase of tepals in both staminate and
pistillate flowers. Aculeate interfloral bracts are found in two species of the African
genus Treculia; the third species of this genus has peltate interfloral bracts (see Berg,
Bull. Jard. Bot. Belg. 47 (1977) 380). Variation of the shape of the processes rather
similar to that as occurs in P. venenosus can be found in some Naucleopsis species.
The staminate and pistillate inflorescences are indistinguishable before anthesis.
KEY TO THE SPECIES
1a. Hairs on the lamina lower surface patent, usually dense; leafy twigs brown tomen-
tose; stipules 0.5 –1 cm long. — Sumatra, Malay Peninsula, Borneo . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. bracteatus
b. Hairs on the lamina lower surface (±) appressed, usually sparse or absent; leafy
twigs whitish to brownish, mostly appressed puberulous; stipules 0.2 – 0.4 cm long.
— Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. venenosus
1. Parartocarpus bracteatus (King) Becc.

Parartocarpus bracteatus (King) Becc., For. Borneo (1902) 632; F.M. Jarrett, J. Arnold Arbor. 41
(1960) 324, t. 21a, d; Corner, Gard. Bull. Singapore 28 (1976) 186, t. 2a; Kochummen, Tree Fl.
Malaya 3 (1978) 164, t. 9; Go, Tree Fl. Sabah & Sarawak 3 (2000) 319. — Artocarpus bracteata
King in Hook.f., Fl. Brit. India 5 (1888) 540; King, Ann. Roy. Bot. Gard. (Calcutta) 2 (1889) 7 p.p.
incl. t. 1B, syncarpium solum; Ridl., Fl. Malay Penins. 3 (1924) 352.
Artocarpus rufescens Kurz, Forest Fl. Burma 2 (1877) 431 (non Miq. 1861).
Tree up to 45 m tall, sometimes with buttresses. Leafy twigs 3 – 8 mm thick, densely

brown (sub)tomentose. Leaves spirally arranged; lamina elliptic to obovate to sub
obovate, 8 – 20(– 28) by 4 –11(–14), coriaceous, apex short-acuminate to acute (to
rounded), base rounded to cordate, margin entire, revolute; upper surface brownish
puberulous, mainly on the main veins; lower surface ± densely brownish puberulous
on the veins; midrib slightly impressed above, lateral veins 8 –18 pairs, tertiary veins
scalariform; petiole 2 – 3.5 cm long, brownish puberulous to subtomentose; stipules
0.5 –1 cm long, brownish puberulous, caducous. Staminate inflorescences axillary, soli-

tary, capitate; peduncle 2 – 3.5 cm long, brown tomentose to puberulous; head globose,
2.5 – 3.5 cm diam.; involucral bracts 3, ovate, 0.5 –1 cm long, brown tomentellous to
puberulous; processes aculeate (or cushion shaped), c. 3 mm long and often slightly
curved (or cushion shaped), (sub)glabrous; stamens 2, up to 8 mm long, filaments free
or connate at the base, anthers 2.5 – 3 mm long, apiculate. Pistillate inflorescences
axillary, solitary, capitate; peduncle 3 – 5.5 cm long, brown tomentose to puberulous;
head globose, 2 – 3.5 cm diam.; involucral bracts 2 – 4, ovate, 0.5 –1 cm long, brown
tomentellous to puberulous; flowers numerous; processes aculeate, 2 – 4 mm long, often
± curved, basally connate in groups of 2 – 4 or free, subglabrous; stigmas subulate, 1–1.5
mm long. Infructescences subglobose, 5 – 9 cm diam., ± lobed, with numerous fruits;
processes conical, up to 8 mm long. — Fig. 22a – c.
Distribution — Peninsular Thailand; in Malesia: Sumatra (also Simalur Island and
Banka), Malay Peninsula, Borneo.
Habitat — Evergreen forest at altitudes up to 700 m.
Note — Two collections from Simalur Island (off Sumatra) match P. bracteatus in
the features of the indumentum and the leaves, but the processes of the inflorescences
are cushion shaped and resemble those of a form of P. venenosus.
2. Parartocarpus venenosus (Zoll. & Moritzi) Becc.

Parartocarpus venenosus (Zoll. & Moritzi) Becc., For. Borneo (1902) 632; F.M. Jarrett, J. Arnold
Arbor. 41 (1960) 326, t. 21b, c, e – h; Backer & Bakh.f., Fl. Java 2 (1965) 18; Kochummen, Tree
Fl. Malaya 3 (1978) 165, t. 9; Go, Tree Fl. Sabah & Sarawak 3 (2000) 323, t. 14.
Artocarpus venenosa Zoll. & Moritzi, Natuur-Geneesk. Arch. Ned.-Indie 2 (1845) 213; Flora 30 (1847)
471; Miq. in Zoll., Syst. Verz. 2 (1854) 89, 95; K. Schum., Notizbl. Königl. Bot. Gart. Berlin 1
(1895) 48. — Gymnartocarpus venenosa (Zoll. & Moritzi) Boerl., Icon. Bogor. 1 (1897) 73, t. 24,
25; Handl. Fl. Ned. Ind. 3 (1900) 371; Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 28.
— Artocarpus venenosa Zoll. & Moritzi var. tylophylla Miq. in Zoll., Syst. Verz. 2 (1854) 89, 95.
— Artocarpus tylophylla (Miq.) Miq., Fl. Ned. Ind. 1, 2 (1859) 289.
Artocarpus cerifera Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 212.
Artocarpus riedelii Miq., Ann. Mus. Bot. Lugduno-Batavi 3 (1867) 213.
Parartocarpus beccarianus Baill., Adansonia 11 (1875) 294; Becc., For. Borneo (1902) 632.
Artocarpus forbesii King in Hook.f., Fl. Brit. India 5 (1888) 539; Ann. Roy. Bot. Gard. (Calcutta) 2
(1889) 7, t. 1A; Ridl., Fl. Malay Penins. 3 (1924) 352; S. Moore, J. Bot. 63, Suppl. (1925) 112.
— Parartocarpus venenosus (Zoll. & Moritzi) Becc. subsp. forbesii (King) F.M. Jarrett, J. Arnold
Arbor. 40 (1959) 9 –12, t. 2a – f, t. 3g – i, t. 4d; 41 (1960) 137, 333; Corner, Gard. Bull. Singapore
28 (1976) 187, t. 2c, 3, 4c; Go, Tree Fl. Sabah & Sarawak 3 (2000) 323.
Artocarpus involucrata K. Schum. in K. Schum. & Hollrung, Fl. Kais. Wilh. Land (1889) 39. — Par
artocarpus involucrata (K. Schum.) K. Schum. & Lauterb., Fl. Schutzgeb. Südsee (1900) 267;
Becc., For. Borneo (1902) 632; Lauterb., Bot. Jahrb. Syst. 62 (1928) 303; Diels, Bot. Jahrb. Syst.
67 (1935) 174; C.T. White, J. Arnold Arbor. 31 (1950) 82.
Parartocarpus papuana Becc., For. Borneo (1902) 633, non S. Moore 1923. — Parartocarpus veneno
sus (Zoll. & Moritzi) Becc. subsp. papuanus (Becc.) F.M. Jarrett, J. Arnold Arbor. 41 (1960) 330;
Corner, Gard. Bull. Singapore 28 (1976) 187, t. 2d, 4a.
Parartocarpus borneensis Becc., For. Borneo (1902) 634. — Parartocarpus venenosus (Zoll. & Mor
itzi) Becc. subsp. borneensis (Becc.) F.M. Jarrett, J. Arnold Arbor. 41 (1960) 332; Corner, Gard.
Bull. Singapore 28 (1976) 186, t. 2c; Go, Tree Fl. Sabah & Sarawak 3 (2000) 323.
Parartocarpus excelsa Becc., For. Borneo (1902) 634.
Artocarpus woodii Merr., Philipp. J. Sci., Bot. 3 (1908) 221; Elmer, Leafl. Philipp. Bot. 2 (1909) 623.
— Parartocarpus woodii (Merr.) Merr., Enum. Philipp. Flow. Pl. 2 (1923) 39. — Gymnartocarpus
woodii (Merr.) Merr., Philipp. J. Sci. 18 (1921) 52; W.H. Br., Bull. Bur. Forest. Philipp. Islands
22, 2 (1921) 270, f. 19.
Gymnartocarpus triandra J.J. Sm., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 233, t. 6 – 8. — Pararto
carpus triandra (J.J. Sm.) J.J. Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1924) 80; F.G. Browne, For.
Trees Sar. & Brunei (1955) 357.
Parartocarpus microcarpus Corner, Gard. Bull. Singapore 28 (1976) 184, t. 1; Go, Tree Fl. Sabah &
Sarawak 3 (2000) 320.
Parartocarpus spinulosus Go, Sandakania 12 (1998) 2, t. 1; Tree Fl. Sabah & Sarawak 3 (2000) 321,
t. 13.
Tree up to 35 m tall, sometimes with low buttresses. Leafy twigs 3 –7 mm thick,

sparsely to rather densely whitish to brown appressedly puberulous. Leaves spirally
arranged; lamina elliptic to obovate to subobovate to narrowly obovate, 4 – 20(– 30) by
2.5 – 8(–10), coriaceous, apex rounded to short-acuminate, base rounded to cuneate or to
subcordate, margin entire, flat or slightly (sometimes distinctly) revolute; upper surface
(sparsely) whitish puberulous, mainly on the main veins to subglabrous, smooth (or
scabrous); lower surface sparsely to rather densely whitish ± appressedly puberulous on
the veins to subglabrous, smooth (or scabrous); midrib slightly impressed to flat above,
lateral veins 6 –16 pairs, tertiary venation scalariform to reticulate; petiole 1.5 – 5.5 cm
long, sparsely to rather densely whitish puberulous; stipules 0.2 – 0.4 cm long, whitish
puberulous, caducous. Staminate inflorescences axillary, solitary, capitate; peduncle
2 – 5.5 cm long, sparsely whitish puberulous; head globose to obovoid, 1.5 – 3 cm diam.;
involucral bracts 3 – 4(– 8), ovate, 0.2 – 0.6 cm long, puberulous or subglabrous; pro
cesses apiculate to conical, umbonate, or ± cushion shaped and depressed in the centre,
0.5 –1.5 mm long, glabrous; stamens (1 or) 2 (or 3), 6 – 8 mm long, filaments partly
connate or free, anthers 1.5 – 2.5 mm long, apiculate or not. Pistillate inflorescences
axillary, solitary, capitate; peduncle 3 –10 cm long, brown tomentose to puberulous;
head globose, 2 – 3.5 cm diam.; involucral bracts 3 – 4(– 8), ovate, 0.2 – 0.6 cm long, pu-
berulous or subglabrous; flowers numerous; processes apiculate to conical, umbonate,
0.5 – 3 mm high, connate in groups of 2 – 4 or free, glabrous or minutely puberulous to
muriculate; stigmas tongue-shaped, 1–1.5 mm long, acute. Infructescences subglobose,
5 –18 cm diam., ± lobed, with numerous fruits; processes conical to aculeate or cushion
shaped and apiculate or depressed in the centre to discoid, up to 10 mm long. — Fig.
22d – j.
Distribution — Peninsular Thailand, Solomon Islands; in Malesia: Sumatra (also
Enggano, Riouw-Lingga Islands), Malay Peninsula, Borneo, Java, Philippines (Luzon,
Mindoro, Samar, Leyte, Mindanao), Celebes, the Moluccas (Aru Islands) New Guinea
(incl. New Britain, Manus Island, Schouten Island, Yapen, Salawati).
Habitat — Forest at altitudes up to 1000 m, in Borneo up to 1800 m; tolerating a
short dry season.
Uses — The latex is used as arrow poison. The ripe infructescence is (said to be)
edible; unripe seeds are poisonous.
Notes — 1. Parartocarpus microcarpus Corner (1976) is included as the morpho-
logical differences between the material referred to this taxon and material referred
to P. venenosus are too small to justify recognition as distinct taxa. In the material
4 cm
2.5 cm
2.5 cm
a
d
f
h
2 mm
8 mm
2 cm
e
g b c i 3 cm
1 cm
7 mm 4 mm
Fig. 22. a – c: Parartocarpus bracteatus (King) Becc. a. Surface view of pistillate inflorescence (arrow
indicating stigmas); b. part of young staminate inflorescence at anthesis in section; c. surface view
of older staminate inflorescence (with stamens exserted between processes). — d – j: Parartocarpus
venenosus (Zoll. & Moritzi) Becc. d. Leafy twig and pistillate inflorescence; e. pistillate inflorescence;
f. surface of pistillate inflorescence; g. in section; h. tangential section of part of pistillate inflorescence
at anthesis; i. infructescence in section; j. surface view of infructescence (arrow indicating stigmas)
(a: Sinclair 39426; b: Corner SFN 28145; c: Beguin 583; d – f: Brass 7476; g – i: Corner SFN 28145;
j: Krukoff 4124).
referred to P. microcarpus the processes of the inflorescences are cushion shaped and
mostly depressed in the centre, a feature which is also found in several collections from
the Philippines and Celebes and from lowland localities. The collections referred to
P. microcarpus are from altitudes between 1000 and 1800 m and may represent a small-
leaved montane form of the species.
2. The species is variable, in particular in the shape of the processes of the inflo-
rescences, and, most pronouncedly so, of the infructescences in which they vary from
discoid to cushion shaped (either with an apiculate or depressed centre) to conical to
aculeate (and then up to 10 mm long). Those with conical or aculeate processes are
similar to those of P. bracteatus.
3. Collections Leopold & Taha SAN 83547 and Soepadmo et al. FRI 41312 (the type
collection of P. spinulosus), both from Sabah, Sandakan district, are aberrant by the
scabrous lamina with a distinctly revolute margin. The processes of the fruiting pistillate
inflorescence are cushion shaped and radially ribbed. Collection De Vogel 6216 from
Celebes has similar processes and the margin of the lamina is also distinctly revolute,
but the surfaces are smooth. Examination of more fertile material is needed before
deciding on the identity and status of these forms.
PRAINEA
Prainea King ex Hook.f., Fl. Brit. India 5 (1888) 546; King, Ann. Roy. Bot. Gard. (Calcutta) 5, 2 (1896)
162, t. 196; Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 1 (1897) 122; Boerl., Handl. Fl. Ned.
Ind. 3 (1900) 337; Becc., For. Borneo (1902) 635; Webbia 5 (1923) 563; Ridl., Fl. Malay Penins.
3 (1924) 358; Steenis, Bull. Jard. Bot. Buitenzorg III, 12 (1932) 259; F.M. Jarrett, J. Arnold Arbor.
40 (1959) 30; Corner, Gard. Bull. Singapore 19 (1962) 243; Kochummen, Tree Fl. Malaya 3 (1978)
165; Go, Tree Fl. Sabah & Sarawak 3 (2000) 326; C.C. Berg, Blumea 50 (2005) 545. — Artocarpus
J. & G. Forst. sect. Prainea (King) Renner, Bot. Jahrb. Syst. 39 (1907) 366.
Trees or climbers, dioecious. Leaves distichous; lamina pinnately veined, entire;

stipules free, lateral. Inflorescences unisexual, solitary or in pairs in the leaf axils, glo-
bose- to obovoid-capitate, pedunculate, with peltate, subpeltate, clavate or spathulate
interfloral bracts. Staminate flowers numerous; perianth tubular, 2-lobed to entire;
stamen 1. Pistillate flowers numerous, free; perianth tubular; ovary free, stigmas 2.
Infructescences with enlarged perianths of some (1– 20) of the flowers, the bracts not
enlarged; pericarp membranaceous; seed with the testa absorbed except the thickened
saucer-shaped basal portion, endosperm absent, embryo straight, longitudinally aligned,
cotyledons equal, appressed faces oblique to median plane of ovary, radicle apical,
minute; interfloral bracts persistent.
Distribution — This Malesian genus comprises two closely related species and rang-
es from the Malay Peninsula to New Guinea.
Morphology — The flowers and bracts of the pistillate inflorescence are free in
contrast to those in the related genera Artocarpus, Hullettia, and Parartocarpus.
In both species, only a small number of the pistillate flowers set fruit. Uncinate hairs
are present in P. scandens.
KEY TO THE SPECIES
1a. Tertiary venation distinctly scalariform; lamina often sparsely hairy above and /or
beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. limpato
b. Tertiary venation reticulate or subscalariform with 1 or 2 ± straight intercostals;
lamina glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. scandens
1. Prainea limpato (Miq.) Beumée ex K. Heyne

Prainea limpato (Miq.) Beumée ex K. Heyne, Nutt. Pl. Ned.-Ind., ed. 2 (1927) 579; F.M. Jarrett,
J. Arnold Arbor. 40 (1959) 34; Kochummen, Tree Fl. Malaya 3 (1978) 167; Go, Tree Fl. Sabah &
Sarawak 3 (2000) 327, t. 15. — Artocarpus limpato Miq., T.S. Liu, Eerste Bijv. (1861) 421; Renner,
Bot. Jahrb. Syst. 39 (1907) 367.
Tree up to 30 m tall, sometimes with buttresses. Leafy twigs 2 – 5 mm thick, sparsely

and minutely puberulous. Leaves distichous; lamina subobovate to elliptic, (4 –)10 –
30(– 40) by (2.5 –)4 –13 cm, often ± asymmetric, at least at the base, coriaceous, apex
acuminate, base cuneate to rounded to subcordate (at one side) or to subauriculate,
margin entire, often ± revolute (towards the base); upper surface glabrous, or puberu-
lous on the veins; lower surface sparsely appressedly puberulous on the main veins
to glabrous, scabridulous or smooth; midrib slightly prominent above, lateral veins
(8 –)12 – 20(– 25) pairs, tertiary venation scalariform; petiole (0.3 –)0.8 – 2(– 3.5) cm
long, sparsely puberulous to subglabrous, the epidermis often flaking off; stipules c. 0.2
cm long, puberulous or glabrous, caducous. Staminate inflorescences axillary, soli-
tary, capitate; peduncle 1.5 – 4(– 6) cm long, minutely puberulous (or glabrous); head
globose, (0.3 –)0.5 –1.5(–1.8) cm diam.; perianth c. 1.5 mm long, densely minutely
puberulous at the apex; stamen c. 1.8 mm long, anther c. 0.4 mm long; bracts peltate,
1.5 – 2 mm long, the apical part 0.2 – 0.3 mm diam., whitish to brownish minutely puber-
ulous. Pistillate inflorescences axillary, solitary, capitate; peduncle 1– 20(– 25) cm long,
minutely puberulous; head globose, 0.8 – 2.5 cm diam.; flowers numerous; perianth 3 – 4
mm long, densely to sparsely minutely puberulous at the apex; stigmas tongue-shaped,
bifid or simple, 0.3 –1 mm long; bracts (sub)peltate, 2.5 – 3 mm long, the upper part
0.3 – 0.4 mm diam., whitish to brownish minutely puberulous. Infructescences sub
globose, 1.5 – 5 cm diam., with numerous protruding fruiting perianths, these ellipsoid
to ovoid, (0.8 –)1.2 – 2.5 cm long, (sub)glabrous, yellowish to reddish at maturity.
Distribution — Sumatra, Malay Peninsula, Borneo, Moluccas, New Guinea.
Habitat — Primary and secondary evergreen forest at altitudes up to 600 m, in New
Guinea up to 1200 m.
Note — The differences between the collections referred to P. limpato and P. pa
puana are rather small and quantitative. Treating the western and eastern Malesian
entities as subspecies appears to be preferable.
a. subsp. limpato
Urostigma diepenhorstii Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 439. — Ficus diepenhorstii (Miq.)
King, Ann. Roy. Bot. Gard. (Calcutta) 1 (1888) 181.
Prainea cuspidata Becc., For. Borneo (1902) 636.
Prainea multinervia Merr., Philipp. J. Sci. 29 (1926) 364; Pl. Elmer. Born. (1929) 46.
Prainea limpato (Miq.) Beumée ex K. Heyne var. longipedunculata Kochummen, Malaysian Forester
41 (1978) 29; Tree Fl. Malaya 3 (1978) 167, t. 10.
Lamina often scabridulous beneath; lateral veins (9 –)13 – 20(– 25) pairs. Staminate
inflorescences: peduncle 2 – 4(– 6) cm long; head 0.8 –1.3(–1.5) cm diameter. Pistillate
inflorescences: peduncle 7– 20(– 25) cm long; head 1– 2(– 2.5) cm diameter. Fruiting
perianth ellipsoid to ovoid, (1.2 –)1.5 – 2(– 2.5) cm long.
Distribution — Sumatra (also Enggano), Malay Peninsula (Kedah), Borneo (also
Bangey Island).
Uses — Ripe infructescences are eaten.
4 cm
2 cm
i 1 mm
2.5 mm
8 mm 8 mm 2 cm
f
1 mm
c d e a b g h
Fig. 23. Prainea limpato (Miq.) Beumée ex K. Heyne subsp. papuana (Becc.) C.C. Berg. a. Leafy twig
with pistillate inflorescences; b. pistillate inflorescence; c. pistillate inflorescence in length section;
d. pistillate flower; e. staminate flower with interfloral bract; f. staminate flower with interfloral bract
in length section; g, h. infructescences; i. fruit in length section: 1 = unexpanded pistillate flower,
2 = fruiting perianth, 3 = embryo, 4 = testa (a, e, f: Beguin 1980; b, c, d, g, i: Beguin 1926; h: Forbes
s.n.).
b. subsp. papuana (Becc.) C.C. Berg

Prainea limpato (Miq.) Beumée ex K. Heyne subsp. papuana (Becc.) C.C. Berg, Blumea 50 (2005)
545. — Prainea papuana Becc., For. Borneo (1902) 635; F.M. Jarrett, J. Arnold Arbor. 40 (1959)
35, f. 1a – e, 3a – c. — Artocarpus papuanus (Becc.) Renner, Bot. Jahrb. Syst. 39 (1907) 367, non
Diels 1936. — Parartocarpus papuana S. Moore, J. Bot 61, Suppl. (1923) 52, non Becc. 1902.
Prainea microcephala J.J. Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1922) 80.
Lamina smooth beneath; lateral veins (8 –)12 –16(– 20) pairs. Staminate inflores
cences: peduncle 1.5 – 4 cm long; head 0.3 – 0.8 cm diameter. Pistillate inflorescences:
peduncle 2 –12 cm long; head (0.5 –)0.8 –1.5(–1.8) cm diameter. Fruiting perianth
subglobose to ovoid, 0.8 –1.3(–1.5) cm long. — Fig. 23.
Distribution — Moluccas (Morotai, Halmahera, Ceram, Ternate, Batjan, Obi), New
Guinea (also Yapen, D’Entrecasteaux and Louisiade Islands, but not in the Bismarck
Archipelago).
Habitat — Rain forest, also in secondary forest at altitudes up to 1200 m.
2. Prainea scandens King ex Hook.f.

Prainea scandens King ex Hook.f., Fl. Brit. India 5 (1888) 547; King, Ann. Roy. Bot. Gard. (Calcutta)
5, 2 (1896) 162, t. 196; Ridl., Fl. Malay Penins. 3 (1924) 358; F.M. Jarrett, J. Arnold Arbor. 40
(1959) 32. — Artocarpus scandens (Hook.f.) Renner, Bot. Jahrb. Syst. 39 (1907) 367.
Prainea frutescens Becc., For. Borneo (1902) 635; F.M. Jarrett, J. Arnold Arbor. 40 (1959) 33; Go,
Tree Fl. Sabah & Sarawak 3 (2000) 327. — Artocarpus frutescens (Becc.) Renner, Bot. Jahrb.
Syst. 39 (1907) 367.
Tree up to 30 m tall or climber. Leafy twigs 1.5 – 2.5 mm thick, sparsely and minutely
puberulous, partly with uncinate hairs, or glabrous. Leaves distichous; lamina sub-
obovate to elliptic, 5 –16 by 3 –7 cm, coriaceous, apex acuminate, base cuneate to
rounded, margin entire; upper and lower surface glabrous; midrib slightly prominent
above, lateral veins 6 –13 pairs, tertiary venation reticulate (to subscalariform with 1 or
2 ± straight intercostals); petiole 0.7–1.8 cm long, sparsely and minutely puberulous
to glabrous, the epidermis often flaking off; stipules c. 0.2 cm long, glabrous, cadu-
cous. Staminate inflorescences axillary, solitary, capitate; peduncle 0.5 –1.5 cm long,
minutely puberulous; head globose to obovoid, 0.4 – 0.6 cm diam.; perianth c. 0.8 mm
long, densely minutely puberulous at the apex; stamen c. 1.2 mm long, anther 0.2 – 0.3
mm long; bracts subpeltate, 0.8 –1 mm long, the apical part 0.2 – 0.5 mm diam., whit-
ish minutely puberulous. Pistillate inflorescences axillary, solitary, capitate; peduncle
1– 2.5 cm long, puberulous; head globose, c. 1 cm diam.; flowers c. 20 – 30; perianth
2.5 – 3 mm long, densely minutely puberulous; stigmas tongue-shaped, 0.5 – 2 mm long;
bracts spathulate to truncate-clavate to subulate or to peltate, 2.5 – 3 mm long, the apical
part 0.2 – 0.5 mm diam., densely whitish minutely puberulous. Infructescences subglo-
bose, 1.5 – 2.5 cm diam., with 1–7 protruding fruiting perianths, these ellipsoid to ovoid,
1.5 – 2.5 cm long, puberulous at the apex, red at maturity. — Fig. 24.
Distribution — Peninsular Thailand; in Malesia: Malay Peninsula (Perak, Selangor),
Borneo.
b
2 mm
2 mm
e
f c 5 mm
3 cm
a
d
2 cm
Fig. 24. Prainea frutescens King ex Hook.f. a. Leafy twig with infructescence; b. pistillate inflores-
cence in length section; c. base of seed with cotyledons and remains of testa; d. infructescence in
length section; e. pistillate flower in length section; f. bract in infructescence (a, c, d, f: SAN 43596;
b, e: Haviland 3102).
Habitat — Forest, sometimes edges of peat swamps; at altitudes up to 800 m.

Notes — 1. The differences between material from the Malay Peninsula and Borneo
are so small that they do not provide a basis to maintain two regional species, P. scan
dens and P. frutescens, respectively.
2. It is not clear how frequently and under which conditions the lianescent habit is
developed.
3. Collection FRI 17082 indicated as ‘Artocarpus sp. A’ by Kochummen (Tree Fl.
Malaya 3 (1978) 134) belongs to this species.
Tribe Castilleae
Castilleae C.C. Berg, Acta Bot. Neerl. 26 (1977) 78. — ‘Olmedieae’ Trécul, Ann. Sci. Nat., Bot.
sér. 3, 8 (1847) 126, excl. Olmedia, being included in Trophis; Corner, Gard. Bull. Singapore 19
(1962) 243.
Trees or shrubs with the architectural model of Cook, dioecious or monoecious,

without uncinate hairs. Leaves alternate, spirally arranged (on the main branches) and
distichous (on the lateral branches); stipules fully to semi-amplexicaul. Inflorescences
on the lateral branches, unisexual, capitate, discoid to cup-shaped with an involucre
of basally attached bracts, pedunculate or sessile, interfloral bracts lacking (? or struc-
tures derived from the perianth). Staminate flowers complete or with reduction in the
perianth and androecium, pistillode mostly absent. Pistillate flowers with the tepals
± connate, ovary usually ± adnate to the perianth, stigmas 2 and equal. Fruit(s) forming
a drupaceous whole with the perianth, with other flowers, or also with the receptacle;
seed large, without endosperm, testa with or without a thickened vascularised part,
embryo longitudinally aligned, with thick, equal or subequal, sometimes unequal coty-
ledons, radicle short and apical.
Distribution — The tribe comprises six neotropical genera (see Berg, Fl. Neotrop.
Monogr. 7 (1972) 1; 83 (2001) 244) for which at present 54 species are recognised. The
tribe is centred in South America. Two monotypic genera are palaeotropical: Antiaris
ranging from the African continent to Madagascar and Yemen and from Sri Lanka to
the Pacific, and Mesogyne confined to tropical Africa (see Berg, Bull. Jard. Bot. Belg.
47 (1977) 323).
Morphology — The tribe is characterised by the tree architecture described as the
model of Cook (Hallé & Oldeman, Essai sur l’architecture et la dynamique de crois
sance des arbres tropicaux (1970) 110). The leaves on the stem and orthotropic branches
are spirally arranged, and in the axils of each of these leaves plagiotropic branches
are formed sylleptically. Inflorescences are borne on these branches, often on axillary
short-shoots. The plagiotropic branches are abscised; the bases of the shed branches are
conical leaving a depression in the stem. This model requires accessory buds which in
the neotropical taxa are lateral but axillary in the palaeotropical genera (see Berg, Acta
Bot. Neerl. 26 (1977) 73).
In the neotropical genera the involucre of imbricate bracts of the pistillate inflo-
rescences covers the outer surface of the receptacle. In the palaeotropical genera the
involucral bracts are more or less scattered on the lower part of the receptacle and
concentrated in the uppermost part.
Chemistry — Cardiotoxic compounds have been found in several representatives of
the tribe (see p. 12 and 17).
KEY TO THE GENERA
1a. Stipules not fully amplexicaul, free; pistillate inflorescences uniflorous; receptacle
of the staminate inflorescence discoid. — Widespread . . . . . . . . . . . . . . Antiaris
b. Stipules fully amplexicaul, fused; pistillate inflorescences with numerous flowers;
receptacle of the staminate inflorescence flabellate and bivalvate (or infundibuli-
form to cyathiform). — Introduced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castilla
CASTILLA
Castilla Sessé in Cerv., Gaz. Lit. Mexico, Suppl. (2.vii.1794) 7; Endl., Gen. Pl. (1837) 282, ‘Castilloa’;
Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 136; Baill., Hist. Pl. 6 (1875) 204; Benth. & Hook.f.,
Gen. Pl. 3 (1880) 372; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 84; Boerl., Handl. Fl.
Ned. Ind. 3 (1900) 328; O.F. Cook, Science n.s. 18 (1903) 436; Pittier, Contr. U.S. Natl. Herb. 13
(1910) 247; Woodson, Ann. Missouri Bot. Gard. 47 (1960) 139; C.C. Berg, Fl. Neotrop. Monogr.
7 (1972) 92; 83 (2001) 269.
Trees, monoecious or (andro)dioecious. Lamina chartaceous, more or less scabrous

above, ± hairy; stipules fully amplexicaul, connate, with distinct parallel veins. Stami
nate inflorescences pedunculate, flabellate and bivalvate (these occurring together with
pistillate ones), infundibuliform to cyathiform, entire to 2-lobed; stamens solitary or
paired along radiating and branching ridges of the receptacle which bears membrana-
ceous, free or connate interstaminal ‘bracts’ (probably representing reduced perianths).
Pistillate inflorescences solitary, sessile, discoid to cupuliform or subglobose, many- to
several-flowered; ovary free, partly adnate to the perianth, or immersed in the recepta-
cle; stigmas strap-shaped, short.
Distribution — The genus is neotropical and comprises three species. Two species
have been economically important for the rubber they yield: the Central American
– Pacific Coastal C. elastica Sessé and the Amazonian C. ulei Warb.
1. Castilla elastica Sessé

Castilla elastica Sessé in Cerv., Gaz. Lit. Mexico, Suppl. (2.vii.1794) 7, Trécul, Ann. Sci. Nat., Bot. sér.
3, 8 (1847) 137, f. 139 –148, ‘Castilloa’; Hook.f., Trans. Linn. Soc. London 2 (1886) 206; Koord.
& Valeton, Bijdr. Boomsoort. Java 11 (1906) 3; Merr., Fl. Manila (1912) 177; Exkurs.-Fl. Java 2
(1912) 97; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 44; Koord., Atlas Baumart. Java (1924) t. 761;
Worth., Ceylon Trees (1959) 418; Woodson, Ann. Missouri Bot. Gard. 47 (1960) 140, f. 50; Little
& Wadsw., Trees Puerto Rico & Virgin Is. 1 (1964) 64, fig.; Backer & Bakh.f., Fl. Java 2 (1965)
19; C.C. Berg, Fl. Neotrop. Monogr. 7 (1972) 94.
Tree up to 30(– 40) m tall, monoecious or (andro)dioecious. Leafy twigs 2 –12 mm

thick, hirsute to hirtellous to velutinous. Leaves distichous on lateral branches; lamina
elliptic to subobovate or to subovate, (10 –)15 – 40(– 55) by (5 –)7– 20(– 30) cm, (almost)
symmetric, chartaceous, apex acuminate, base (sub)cordate, margin dent(icul)ate; up-
per surface hispidulous, ± scabrous; lower surface hirtellous to subhirsute or strigose to
subsericeous, midrib slightly prominent, lateral veins (10 –)18 – 21(– 25) pairs, tertiary
venation scalariform; petiole 0.5 – 2(–10) cm long, hirtellous; stipules 1–11.5 cm long,
sericeous to hirtellous to hirsute, on the margins tomentellous. Staminate inflorescences,
if not accompanying pistillate ones, then peduncle 3 –15 mm long; receptacle flabel-
late and 2- (or 3-)valved, 1– 2.5 by 0.7– 2(– 2.5) cm; involucral bracts in 10 –12 rows;
if accompanying pistillate inflorescences, then peduncle 0 –10 mm long; receptacle
infundibuliform to cyathiform, 5 – 20 mm diam.; involucral bracts in 7–12 rows. Pistil
late inflorescences discoid-capitate, 10 – 20 mm diam., (sub)sessile; involucral bracts in
5 –10 rows, flowers numerous; stigmas strap-shaped. Fruiting perianth red to orange;
fruit 8 –10 by 5 –7 mm.
Subdivision — Three subspecies are recognised, only the typical subspecies intro-
duced in Malesia.
a. subsp. elastica
Ficus gummifera Bertol., Mem. Reale Accad. Sci. Ist. Bologna, ser. 1, 10 (1858) 40, t. 9. — Castill(o)a
gummifera (Bertol.) Standl., Contr. U.S. Natl. Herb. 20 (1917) 34.
Castill(o)a lactiflua O.F. Cook, Science n.s. 18 (1903) 269.
Castill(o)a nicoyensis O.F. Cook, Science n.s. 18 (1903) 438.
Castill(o)a guatemalensis Pittier, Contr. U.S. Natl. Herb. 13 (1910) 34.
Lower surface of the lamina with patent hairs on the lateral veins or also on the
midrib; fruiting perianths only basally connate.
Distribution — Central America, from Mexico to Costa Rica; introduced in Malesia,
mainly in botanical gardens.
Uses — In Java, Sumatra, and the Philippines (still?) used as shade tree in cacao and
coffee plantations.
Note — From this particular subspecies (known as Caucho, Central American Rub-
ber or Panama Rubber) rubber has been extracted commercially and introduced to
many tropical countries, but was soon eclipsed by the commercial success of Hevea
brasiliensis.
ANTIARIS
Antiaris Lesch., Ann. Mus. Hist. Nat. Paris 16 (1810) 478, nom. cons.; R.Br. in Flinders, Voy. Terra
Austral. (1814) 602; Blume, Rumphia 1 (1836) 56; Endl., Gen. Pl. (1837) 280; Trécul, Ann. Sci.
Nat., Bot. sér. 3, 8 (1847) 142, f. 158 –162; Baill., Hist. Pl. 6 (1875) 203; Benth. & Hook.f., Gen. Pl.
3 (1880) 371; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888) 85; Bot. Jahrb. Syst. 33 (1902)
119; Welt Afr. 3, 1 (1915) 33, f. 20; Hutch., Fl. Trop. Afr. 6, 2 (1917) 223; Leandri, Fl. Madagasc.
fam. 55 (1952) 30, f. vi; Corner, Gard. Bull. Singapore 19 (1962) 244; C.C. Berg, Bull. Jard. Bot.
Belg. 47 (1977) 308; 48 (1978) 466.
Ipo Pers., Syn. 2 (1807) 566, nom. rejic.
Antscha Horsf., Verh. Batav. Gen. 7 (1814) 8.
Toxicaria Aepnel. in Steud., Nomencl. ed. 2 (1841) 694, non Schreb. 1783.
Lepurandra Graham, Cat. Pl. Bombay (1839) 193.
Trees, monoecious or dioecious. Leaves chartaceous to subcoriaceous, more or less

hairy, margin subentire to denticulate; stipules semi-amplexicaul, free. Inflorescences
on short-shoots in the axils of the leaves or below the leaves, with 1 or 2 pistillate in-
florescences (on the lower nodes of the short-shoot) and/or up to c. 8 staminate ones.
Staminate inflorescences discoid, pedunculate, bracts many, basally attached, those
on the margin of the receptacle in 1– 3 rows, imbricate, forming an involucre; flowers
many, with 2 –7 free tepals; stamens 2 – 4, straight before anthesis, anthers latrorse to
extrorse; pistillode absent. Pistillate inflorescences sessile or pedunculate, with an in-
volucre of several basally attached imbricate bracts; flowers 1, occasionally 2; perianth
4-lobed, the lower part adnate to the receptacle; ovary adnate to the embedded part of
the perianth; stigmas 2. Infructescences with an orange to scarlet, fleshy receptacle,
forming a whole with the fruit, the involucral bracts scattered on its outer surface; en-
docarp crustaceous.
Distribution — The genus comprises a single very variable species, occurring in
continental tropical Africa, Yemen, Madagascar, and from Sri Lanka to Tonga.
1. Antiaris toxicaria Lesch.

Antiaris toxicaria Lesch., Ann. Mus. Hist. Nat. Paris 16 (1810) 478, t. 22; Blume, Rumphia 1 (1836)
56, t. 22, 23; Hook., Companion Bot. Mag. 1 (1836) 310, t. 17, cum syn. Antiaris dubia Span.,
nomen; Benn. in Benn. & R.Br., Pl. Jav. Rar. (1838) 52, t. 13; Trécul, Ann. Sci. Nat., Bot. sér. 3, 8
(1847) 143; Miq., Fl. Ned. Ind. 1, 2 (1859) 291; Fl. Ned. Ind., Eerste Bijv. (1861) 423; Ann. Mus.
Bot. Lugduno-Batavi 3 (1867) 214; Baill., Hist. Pl. 6 (1875) 203, f. 119 –121; Kurz, Forest Fl.
Burma 2 (1877) 462; Fern.-Vill., Nov. App. (1880) 102; S. Vidal, Sin. Gen. Pl. Leños Filip. (1883)
40, t. 88 f. A; Hook.f., Fl. Brit. India 5 (1888) 537; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1
(1888) 85, f. 64; Hook.f. in Trimen, Handb. Fl. Ceylon 4 (1898) 97; Becc., For. Borneo (1902) 594;
Koord. & Valeton, Bijdr. Boomsoort. Java 11 (1906) 274; Renner, Bot. Jahrb. Syst. 39 (1907) 374;
Merr., Philipp. J. Sci., Bot. 2 (1907) 111; Koord., Exkurs.-Fl. Java 2 (1912) 97; Merr., Interpr. Herb.
Amboin. (1917) 192; Enum. Born. Pl. (1921) 220; Enum. Philipp. Flow. Pl. 2 (1923) 44; Ridl., Fl.
Malay Penins. 3 (1924) 351; Koord., Atlas Baumart. Java (1924) t. 762; Gagnep., Fl. Indo-Chine 5
(1928) 721, f. 98: 1– 8; Corner, Wayside Trees Malaya (1940) 648; Steenis, Blumea 6 (1948) 258;
Worth., Ceylon Trees (1959) pl. 419; Corner, Gard. Bull. Singapore 19 (1962) 248; C.C. Berg, Bull.
Jard. Bot. Belg. 47 (1977) 310; 48 (1978) 466; Kochummen, Tree Fl. Malaya 3 (1978) 120; Boer,
Brink & Sosef, Prosea 12, 1 (1999) 126, cum t.; Go, Tree Fl. Sabah & Sarawak 3 (2000) 185, t. 1.
— [Arbor toxicaria mas Rumph., Herb. Amboin. 2 (1741) 263, t. 87.]
Trees up to 50 m tall, (always?) deciduous. Leafy twigs 2 – 5 mm thick, brownish

hirtellous to puberulous. Leaves distichous on the lateral branches; lamina elliptic to
(sub)obovate or to (sub)ovate, 5 – 20(– 30) by 2.5 –10(–12) cm, ± asymmetric, charta-
ceous to subcoriaceous, apex short-acuminate to subacute or obtuse (to rounded), base
cordate to rounded, margin entire or denticulate (towards the apex); upper surface pu-
berulous to hispidulous, scabrous or (sub)glabrous and smooth; lower surface densely to
(very) sparsely puberulous to hirtellous or to subtomentose on the veins, smooth; midrib
slightly prominent to flat above, lateral veins 7–12 pairs, tertiary venation scalariform;
petiole 0.3 –1.2 cm long, puberulous to hirtellous; stipules 0.4 –1 cm long, puberulous,
caducous. Staminate inflorescences up to 4 together, in leaf axils, on short-shoots or
at the base of lateral branches; peduncle 5 –15 mm long, puberulous; head discoid,
5 –12(– 20) mm diam.; involucral bracts in c. 4 rows, ovate, 1– 2 mm long, minutely
puberulous; tepals (2 –)3 – 5(–7), spathulate-cucullate, 1– 2 mm; stamens 3 or 4, 1– 2
mm long, anthers 0.8 –1.8 mm long. Pistillate inflorescences solitary or in pairs on
short-shoots; peduncle 3 – 5 mm, in fruit up to 1 cm long, puberulous; receptacle with
6 –12 bracts, most of them crowded at the apex, 0.5 –1 mm long, minutely puberulous;
stigmas 5 –15 mm long. Infructescences ellipsoid, 1.2 – 4.5 cm long, red to crimson at
maturity; seed ellipsoid, 0.5 – 2 cm long. — Fig. 25.
Distribution — Tropical Africa, Madagascar, Yemen, from Sri Lanka and South India
to South China, Indochina, Thailand, and the Pacific (Tonga); in Malesia: throughout.
Uses — The milk sap is used to prepare arrow poison, the active substances in which
cardiac glycosides (cardenolides), including forms of antiarin, which arrest the heart
(see Bennett in Bennett & R. Brown, Pl. Jav. Rar (1838) 52; Bissett, Ann. Bogor. 2
(1957) 219, Planta Med. 10 (1962) 143; Lloydia 29 (1966)1, 172; Burkill, Dict. (1935)
174; K. Heyne, Nutt. Pl. (1950) 566; Teijsm., Natuurk. Tijdschr. Ned.-Indië 11 (1856)
175; Wehrli, Schindler & Reichstein, Helvet. Chim. Acta 45 (1962) 1083; Wehrli, Hel-
vet. Chim. Acta 45 (1962) 1206; Boer, Brink & Sosef, Prosea 12, 1 (1999) 126). The
concentration of cardiac glycosides varies and may be too low for the preparation of
2 cm
3 mm
g f
h
4.5 mm
7 mm
e
4 mm
1 cm
Fig. 25. Antiaris toxicaria Lesch. a. Seedling; b. pistil-

late inflorescence, one-flowered, at anthesis; c. base
of the pistillate inflorescence at anthesis; d, e. pis-
b tillate inflorescence at anthesis: d. in length section,
e. in cross section; f. staminate inflorescence at an-
thesis in lenght section; g. mature stamen; h. tepals of
c staminate flower (a: FRI bb 1379, b – h: material from
1 cm 1 cm Singapore Botanic Gardens).
arrow poison. Outside western and central Malesia there seems to be no native knowl-
edge that the latex is poisonous. Latex of some species of Naucleopsis (a neotropical
genus of the tribe Castilleae) is used to prepare arrow poison in parts of South America
(see Bisset & Hylands, Econ. Bot. 20 (1977) 237).
Notes — 1. The species is very variable. Several subspecies and varieties have been
recognised: for the African continent three subspecies, for Madagascar two, and for the
Asian – Australasian region two (see Berg, Bull. Jard. Bot. Belg. 47 (1977) 309). Juve-
nile material of all these subspecies is (rather) similar, but adult (and fertile) material
can be distinguished reasonably well. The boundary between the two Asian subspecies
is not clear in eastern Malesia (Moluccas and Lesser Sunda Islands).
2. Ranging from West Africa to Polynesia, this species is the most widespread of the
Moraceae.
a. subsp. toxicaria
Ipo toxicaria Pers., Syn. Pl. 2 (1807) 566.
Antiaris innoxia Blume, Rumphia 1 (1837) 172, t. 54; Miq., Fl. Ned. Ind. 1, 2 (1859) 292.
Lepurandra saccidora Nimmo in Graham, Cat. Pl. Bombay (1839) 193. — Antiaris saccidora (Nimmo)
Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 3 (1851) 232; Wight, Icon. Pl. Ind. Orient. (1853)
t. 1958.
Antiaris rufa Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 423; Koord., Versl. Minahasa (1898) 592.
Antiaris ? palembanica Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 424.
Antiaris zeylanica Seem., Bonplandia 10 (1862) 4.
Leafy twigs appressedly to patently puberulous to hirtellous; periderm usually per-

sistent. Lamina elliptic to (sub)obovate, apex acuminate to rounded; upper surface
hirtellous to hispidulous to puberulous, scabrous or (in adult material) to smooth; lower
surface appressedly to patently puberulous to hirtellous. Infructescences 1.2 – 2.5 cm
long.
Distribution — India (southern & Andaman Islands) Sri Lanka, South China (incl.
Hainan), Indochina, Myanmar, Thailand; in Malesia: Sumatra, Malay Peninsula, Bor-
neo, Java, Philippines (Palawan, Luzon, Mindoro, Guimares), Celebes, Lesser Sunda
Islands (Sumba, Flores, Alor), Moluccas (?).
Habitat — Forest at altitudes up to 1500 m.
b. subsp. macrophylla (R.Br.) C.C. Berg

Antiaris toxicaria Lesch. subsp. macrophylla (R.Br.) C.C. Berg, Bull. Jard. Bot. Belg. 47 (1977) 309.
— Antiaris macrophylla R.Br. in Flinders, Voy. Terra Austral. (1814) 602, t. 5; Benth., Fl. Australia
6 (1873) 179. — Antiaris toxicaria Lesch. var. macrophylla (R.Br.) Corner, Gard. Bull. Singapore
19 (1962) 248; Backer & Bakh.f., Fl. Java 2 (1965) 20. — Antiaris bennettii Seem., Bonplandia 9
(1861) 259; 10 (1862) 3, t. 7; Fl. Vit. (1868) 253, t. 72.
Leafy twigs sparsely appressedly (to patently) puberulous; periderm usually flaking
off. Lamina elliptic to subovate, apex acuminate to subacute to obtuse, upper surface
(sparsely) puberulous (mainly in the midrib), smooth (in juvenile material smooth or
scabridulous); lower surface sparsely appressedly puberulous to subtomentose on the
veins. Infructescences 2 – 4.5 cm long.
Distribution — Pacific (Solomon Islands, New Hebrides, Vanuatu, Fiji, Tonga), Aus-
tralia (Arnhemland, Company’s Island); in Malesia: Moluccas (Batjan?, Halmahera,
Aru Islands), Lesser Sunda Islands (only Timor, Wetar, and Tanimbar Islands?), New
Guinea (incl. Bismarck Archipelago).
Habitat — Forest at altitudes up to 800 m.
Uses — The infructescences are edible and the (sub)species has been carried to Fiji,
Tonga, and Company’s Island by sea farers (see A.C. Smith, Fl. Vitensis Nova 2 (1981)
200). The small size of the plants found in Fiji (either shrubs or small trees) is linked
to this introduction. It should be noted that specimens of A. toxicaria subsp. humbertii

(Leandri) C.C. Berg of Madagascar are shrubs or small trees and the same habit oc-
curs in the other Madagascan subspecies (see Berg, Bull. Jard. Bot. Belg. 47 (1977)
318 – 321).
Tribe Dorstenieae
Dorstenieae Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 510; Bureau in A.DC., Prodr. 17 (1873) 254;
Benth. & Hook.f., Gen. Pl. 3 (1880) 346; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888)
79; Corner, Gard. Bull. Singapore 19 (1962) 250; C.C. Berg, Proc. Kon. Ned. Akad. Wetensch. C,
91 (1998) 359.
Brosimeae Trécul, Ann. Sci. Nat., Bot. sér. 3, 8 (1847) 146; Corner, Gard. Bull. Singapore 19 (1962)
250; C.C. Berg, Fl. Neotrop. Monogr. 7 (1972) 161.
Trees, shrubs (of various growth habit) or in Dorstenia herbs, mostly monoecious,
less commonly androdioecious or dioecious, usually with uncinate hairs. Leaves dis-
tichously or spirally arranged; lamina pinnately, subtriplinerved, (sub)palmately, or
radially veined; stipules large to small, fully amplexicaul to lateral. Inflorescences
bisexual (or secondarily unisexual), discoid to turbinate or to cup-shaped, circular or
shaped otherwise, never distinctly involucrate, interfloral bracts mostly present and
peltate. Staminate flowers often with reductions in the perianth (down to absent) and
the stamens (down to one), pistillode mostly absent. Pistillate flowers mostly connate
with other flowers and/or largely embedded in the receptacle; stigmas 2 and equal, but
in Dorstenia often unequally long or only 1. Fruit a dehiscent drupe(let) or forming
a drupaceous whole with other flowers and/or the receptacle; seed large and without
endosperm (in Dorstenia small and with endosperm), testa sometimes with a thickened
vascularised part; embryo with thick and often unequal cotyledons, or if seeds small,
then with flat and equal cotyledons.
Distribution — This tribe comprises eight genera with in total 87 species. It has a
distinct West Gondwanan distribution. Three genera of woody plants with in total 18
species are confined to the Neotropics and concentrated in the South American conti-
nent. Four genera of woody plants with in total 5 species are endemic to the African
region. The eighth genus, Dorstenia, is largely herbaceous and is represented in both
major flora regions. Only one of the 105 species, Dorstenia indica Wight, is Asian (Sri
Lanka and W India).
Morphology — The tribe is characterised by bisexual inflorescences with one or
more pistillate flowers in the centre and the staminate flowers mostly peripheral. Peltate
bracts and uncinate hairs are common.
DORSTENIA
Dorstenia L., Sp. Pl. (1753) 121; L., Gen. Pl. ed. 5 (1754) 56; Endl., Gen. Pl. (1837) 278; Bureau in
A.DC., Prodr. 17 (1873) 258; Baill., Hist. Pl. 6 (1875) 196; Benth. & Hook.f., Gen. Pl. 3 (1880)
366; Hook.f., Fl. Brit. India 5 (1888) 494; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3, 1 (1888)
79; Rendle, Fl. Trop. Afr. 6, 2 (1916) 25; Leandri, Mem. Inst. Sci. Madagascar, Sér. B, Biol. Vég.
1 (1948) 29; Corner, Gard. Bull. Singapore 19 (1962) 250 (passim); C.C. Berg & Hijman, Ilicifolia
2 (1999) 1; C.C. Berg, Fl. Neotrop. Monogr. 83 (2001) 144.
Kosaria Forssk., Fl. Aegypt.-Arab. (1775) 164.

Sychinium Desv., Mém. Soc. Linn. Paris 4 (1826) 216, ‘Sichinum’.
Ctenocladus Engl., Bot. Jahrb. Syst. 57 (1921) 246.
Craterogyne Lanj., Recueil Trav. Bot. Neerl. 32 (1935) 272.
Herbs, often partly ± succulent, rhizomatous or tuberous, or shrubs. Leaves spirally

arranged or distichous; lamina pinnately, less often palmately or radially veined; stip-
ules lateral, free. Inflorescences bisexual, in the leaf-axils, discoid to turbinate, in out-
line circular, elliptic, ± stellate or navicular, receptacle outside bracteate or mostly with
marginal and/or submarginal appendages; flowers connate. Staminate flowers numer-
ous; tepals (1 or) 2 or 3 (or 4), free or basally connate; stamens 2 or 3; pistillode usually
absent. Pistillate flowers in the central part of the inflorescence, 1– numerous; perianth
tubular, with only the apex free from the surrounding flowers; ovary free; stigmas 1 or
2, filiform to band-shaped, equal or unequal. Fruit a dehiscent drupe, the white fleshy
part pushing out the endocarp body (pyrene) if large or ejecting the endocarp body if
small; endocarp body (especially if large) subglobose and smooth or tetrahedral and
tuberculate; seed large and without endosperm or small and with endosperm, cotyledons
thick and unequal or flat and equal.
Distribution — The genus comprises 105 species, 46 in the Neotropics, 1 in Asia (Sri
Lanka, W India), 58 in continental Africa extending to Madagascar, the Arabian Penin-
sula, and Socotra. Dorstenia contrajerva is introduced in several tropical countries.
Morphology — The majority of the species are herbaceous, eleven African species
are woody (shrubs or even treelets) and some in Africa and Brazil are suffrutescent.
1. Dorstenia contrajerva L.
Dorstenia contrajerva L., Sp. Pl. (1753) 121; F. Overbeck, Jahrb. Wiss. Bot. 66 (1924) 467; Wood-
son, Ann. Missouri Bot. Gard. 47 (1960) 177, f. 62; Backer & Bakh.f., Fl. Java 2 (1965) 17; C.C.
Berg, Fl. Neotrop. Monogr. 83 (2001) 198. — Dorstenia quadrangularis Stokes, Bot. Mat. Med.
4 (1812) 338.
Dorstenia contrajerva L. var. houstonii L., Sp. Pl. (1753) 121. — Dorstenia houstonii (L.) L., Sp. Pl.
ed. 2. (1762 –1763) 176. — Dorstenia quadrangularis Stokes var. integrifolia Stokes, Bot. Mat.
Med. 4 (1812) 339.
Dorstenia alexiteria L., Syst. Nat. ed. 10. (1759) 889.
Dorstenia quadrangularis Stokes var. pinnatifida Stokes, Bot. Mat. Med. 4 (1812) 341.
Dorstenia quadrangularis Stokes var. sinuata Stokes, Bot. Mat. Med. 4 (1812) 339.
Dorstenia palmata Willd. ex Schult., Mant. 3 (1827) 317, non Engler (1894).
Dorstenia maculata Lem., Ill. Hort. 10 (1863) t. 362. — Dorstenia contrajerva L. var. maculata (Lem.)
Bureau in DC., Prodr. 17 (1873) 260.
Dorstenia contrajerva L. subsp. tenuiloba Blake, Contr. U.S. Natl. Herb. 24 (1922) 2, t. 1. — Dorstenia
contrajerva L. var. tenuiloba (Blake) Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23
(1944) 44.
Herb, usually rosulate; internodes usually short. Leaves spirally arranged; lamina
broadly ovate to cordiform to subhastate, pinnately to subpalmately to subpedately
lobed to parted or (almost) entire, 4 – 25 by 3.5 – 30 cm, membranaceous, apex acute to
subacuminate, base cordate to acute, margin crenate-dentate (to entire); upper surface
hirtellous to subhirsute to strig(ill)ose, usually ± scabrous; lower surface puberulous to
hispidulous, often ± scabrous; lateral veins 3 –7 pairs; petiole 3 – 20 cm long; stipules
0.2 – 0.6 cm long, puberulous. Inflorescences green or the margin of the receptacle
purplish; peduncle 5 – 30 cm; receptacle (0.5 –)1.5 – 3 cm wide, ± quadrangular and /or
± irregularly lobed, excentrally attached.
Distribution — Mexico to Panama, West Indies, Colombia, northern Venezuela, west
ern Ecuador, northern Amazonian Peru; in Malesia: introduced (as ornamental): Java.
Tribe Ficeae
See Moraceae: Ficus, Flora Malesiana, Series I, Volume 17, Part 2 (2005).
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Flora Malesiana. Series I: Seed Plants

Article in Taxon · February 2006

The user has requested enhancement of the downloaded file.

The following treatments of Artocarpus, Hullettia, Parartocarpus, and Prainea are

Flowers — Moraceous flowers are small, monochlamydeous, and usually unisexual.

PHYTOCHEMISTRY AND CHEMOTAXONOMY

These features are: production of furanocoumarins, isoflavones (osajin, pomiferin),

1) This paragraph is an addition by C.C. Berg.

Pollination — Wind-pollination occurs in Moraceae with urticaceous type stamens and

DELIMITATION AND SUBDIVISION OF THE FAMILY

spp., neotropical and characterized by simply constructed, Moreae-like inflorescences

KEY TO THE TRIBES

KEY TO THE GENERA

1a. Plants herbaceous or suffrutescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

7a. Stipules connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castilla

22a. Inflorescences spicate, all flowers sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Trees, shrubs, or in Fatoua, herbs or subshrubs, dioecious or sometimes monoecious,

continent. The number of species is c. 70, c. 47 of them occur in the Asian–Australasian

Staminate flowers and pollination — This tribe is characterised by stamens similar

Inflorescences — The inflorescences of the Moreae are mostly unisexual. Bisexual

Fruits — Dehiscent drupes occur in Bleekrodea, Broussonetia, Fatoua, and some of

KEY TO THE GENERA OF THE MOREAE

1a. Plants herbaceous (or suffrutescent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fatoua

KEY TO THE SPECIES

Tree up to 10(–15) m tall. Leafy twigs 1– 4 mm thick, (minutely and/or sparsely)

2. Morus macroura Miq.

Tree up to 32 m tall, deciduous. Leafy twigs 1.5 – 3 mm thick, puberulous to hirtel-

Fig. 1. Morus macroura Miq. a. Leafy twig with axil-

Map 1. Distribution of Morus macroura Miq.

densely puberulous to hirtellous, caducous (or subpersistent). Staminate inflorescences

Shrubs or treelets, monoecious, unarmed. Leaves distichous; lamina pinnately veined;

subcapitate, sometimes subinvolucrate. Staminate flowers several to one (or lacking),

1. Bleekrodea insignis Blume

lamina pinnately veined; stipules free, semi-amplexicaul. Inflorescences unisexual, usu-

KEY TO THE SPECIES

1a. Leaves spirally arranged or subopposite; lamina scabrous above. — Widespread,

1. Broussonetia kurzii (Hook.f.) Corner

Climber, deciduous. Leafy twigs 1– 3 mm thick, appressedly puberulous. Leaves

2. Broussonetia luzonica (Blanco) Bureau

Fig. 3. Broussonetia luzonica (Blanco) Bureau.

3. Broussonetia papyrifera (L.) Vent.

Tree up to 35 m tall, deciduous. Leafy twigs 1.5 – 3 mm thick, subtomentose to hirtel-

ral bracts clavate to subpeltate, 1–1.5 mm long, densely puberulous to tomentellous.

Subshrubs or herbs, monoecious, latex absent. Leaves distichous; lamina pinnately

1. Fatoua villosa (Murray) Nakai

subpersistent. Inflorescences subcapitate, 0.3 –1 cm diam.; peduncle 0.5 – 2.5 cm long,

short-shoots, often yellow (dye-containing) glands embedded in the bracts and/or

Climbers or shrubs. Staminate inflorescences globose-capitate; stamens straight in

1. Maclura cochinchinensis (Lour.) Corner

Shrub, often much-branched with long scrambling sprays, or climber, with up to

puberulous to tomentellous; head globose, 0.4 –1 cm diam.; tepals c. 1.5 mm long,

Trees or shrubs, dioecious or monoecious, unarmed or armed with thorns, uncinate

Relationships — The genus shows parallel morphological differentiation to the ge-

KEY TO THE SECTIONS

Albrandia Gaudich. in Freyc., Voy. Uranie, Bot. (1830) 509.

Trees or shrubs, monoecious or dioecious, without spines. Leaves distichous; lamina

KEY TO THE SPECIES

1. Streblus asper Lour.

Fig. 8. Streblus glaber (Merr.) Corner subsp. uro

Fruits — Ellipsoid (to globose), with a ± crustaceous to cartilaginous pericarp or endo