Manten 66 Half

Earth-Science Reviews - Elsevier Publishing C o m p a n y , A m s t e r d a m - Printed in The N e t h e r l a n d s
HALF A CENTURY OF MODERN PALYNOLOGY
A. A. MANTEN
Secretary and Editor, Second International Conference on Palynohz~y,

Cortezlaan 9, Utrecht (The Netherlamts)
SUMMARY
A short review of the history of palynology is presented. This branch of science

received its greatest impetus from the introduction of the pollen diagram, 50
years ago. Emphasis is laid on the development of palaeopalynology, but the
history of pollen morphology, medical pollen studies and melittopalynology
are also briefly discussed, as well as work on chitinozoans and fossil micro-
plankton of botanical affinity. International palynological meetings are also
reported.
INTRODUCTION
In 1966, palynology celebrates its 50th anniversary. Modern palynology is con-

sidered to have come into existence at the Sixteenth Scandinavian Meeting of
Natural Scientists, in Christiania (now Oslo), in 1916, where the Swede Lennart
yon Post presented the first pollen diagrams. This started the extensive appli-
cation of pollen-and-spore studies to various geological, archaeological and
palaeobotanical purposes.
Before 1916, however, various kinds of other pollen researches had al-
ready been carried out by botanists, physicians and some geologists. Therefore,
the year 1916 was not an absolute starting point. But it was the geological interest
in fossil pollen that brought the largest group of pollen students to the family
of modern palynologists and that had a catalyzing effect also on the activities
of other groups, notably the pollen morphologists. So at least, 1916 was a most
significant year in the development of palynology.
The semi-centennial is a good moment to focus some extra attention on
palynology. It is interesting to look back for a while at the history of this branch
of science and also to pay attention, in a second paper (MANTEN, 1966b), to a
number of current trends within palynology. Both contributions will serve to
show the extensive and still growing value of this field for a variety of other areas
of scientific activity.
Earth-Sci. Rev., 2 (1966) 277-316

278 A . A . MANTEN
This paper will, thus, concentrate on a short historical survey of the devel-
opment of palynology. It should be pointed out that the purpose of such a survey
is to show how palynology after half a century has reached its present position.
This paper is certainly not a complete review article. Not only would there not
be sufficient space for such an attempt, but also the general historical picture
would tend to be obscured if too many examples and data were presented. I am
sure that every specialist will find that much important work is omitted from
this survey and what is mentioned is partly a personal, call it arbitrary, selection.
I admit this and, where required, I do apologize. Nevertheless, I hope that the
paper will serve to show the various stages of gestation, youth and puberty that
this branch of science has undergone to reach the early maturity it has today.
For the convenience of describing the history of palynology, the boundaries
between the successive stages are set at the years 1916 (first pollen diagrams)
and 1944 (coining of the name palynology).
POLLEN STUDIES BEFORE 1 9 1 6
Pollen morphology
The first studies on pollen were evidently done by botanists. These started in
the middle of the seventeenth century, as a result of an improvement in the
construction of microscopes. Soon after, the Englishman Nehemiah Grew made
the significant discovery that pollen grains in different plant species are of different
size and form, but that the pollen grains of flowers belonging to the same species
are all alike.
In the seventeen-sixties, notable contributions to pollen morphology were
made by J. G. KiSlreuter, who, despite the fact that his microscopes were
only very imperfect, discovered that the outer covering of pollen grains consists
of two distinct coats. In some species, he also noted the spines and sculpturing
on the outer coat of the pollen grain and the elasticity of that coat.
In the years 1790-1840, Francis Bauer, working in the Royal Botanic Gar-
dens at Kew, Great Britain, made very accurate drawings of many pollen grains.
However, the majority of his work was never published and consequently, he
had only very little influence on the development of pollen science.
The first successful use of pollen characters in classification appears to
have been that of John Lindley, in 1830, in genera and species of orchidaceous
plants.
Hugo YON MOHL, of German descent, but professor of physiology at Bern,
Switzerland, published in 1834 a work entitled Ober den Bau und die Formen
yon Pollenk6rner, which was a major contribution to knowledge of the structure
Earth-Sci. Rev., 2 (1966) 277-316

HALF A CENTURY OF MODERN PALYNOLOGY 279
of pollen grains and the terminology needed to describe it. The greater part of
the work gives a detailed descriptive classification of pollen forms.
Other significant contributions came at about that time and also later
from Carl Julius FRITZSCHE, who in his later years, however, left botany to
become one of the great chemists of his time. Fritzsche's pollen observations
and drawings were often more accurate than those by Von Mohl, but he was
less interested in drawing conclusions from his work.
In 1889, Carl Albert Hugo FISCHER obtained his doctor degree with a
thesis entitled Beitrdge zur vergleichenden Morphologie der Pollenkrrner. This
thesis was based on two questions: (1) how is the outer layer of a pollen grain
formed? and (2) in what way do plants which are related in their outward form
agree in their pollen-grain structures? To find answers to these questions, he
studied thoroughly pollen grains of over 2,000 plant species from 158 families.
His studies were the first to greatly benefit from the introduction, in 1884, of the
apochromatic lens. Fischer's thesis could have been a most promising start to
a pollen-morphological career, but unfortunately it was also the end of it, as he
was later more interested in colloid chemistry, plant nutrition, soil chemistry and
applied botany.
Medical pollen studies
A specialized line of pollen studies developed in medicine, in relation to the

occurrence of inhalent allergic diseases, such as hay fever and hay asthma.
The first scientific paper on hay fever appeared in 1819. It was written by
the Englishman John BOSTOCK, who presented his own case. A great variety of
opinion about the causes and nature of hay fever was expressed, until BLACKLEY
(1873) carried out the first day-to-day studies of pollen incidence, using horizon-
tal slides, mainly. He demonstrated that the disease is due to exposure to the pollen
from certain plants. His work only received scant attention, as physicians had
already recognized that a sea voyage or a sojourn at the seashore often gave
relief, and Blackley's theory offered nothing new in treatment. Apart from an
isolated attempt by Lockwood, in the U.S.A., no further aerobiological research
was carried out until 1916, when William Scheppegrell began his local sampling
and counting in New Orleans. He revived the use of horizontal slides to trap
pollen, the so-called "gravity slide" method (SCHEPPEGRELL,1923).
A number of years before Scheppegrell started his pollen studies, light
had also been thrown on the nature of hay fever. The Germans Weichardt, in
1905, and Wolff-Eisner, in 1906, were the first to suggest, and the Englishman
Noon, in 1911, was the fiirst to prove that the disease is an anaphylactic pheno-
menon. This implies the destruction of a normal or natural immunity factor
during an incubation period.
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280 A. A. MANTEN
Observations of fossil pollen
Fossil pollen grains were only observed for the first time in February 1836. This
was by H. R. G/3ppert, who studied the Miocene browncoal of Salzhausen in
Hessen, Germany (G6PPERT, 1836). The specific identification of seeds and other
small fossils found in the browncoal required some strong magnification, and
the use of a microscope naturally led to the discovery of even smaller fossils,
including pollen grains.
The conservation of pollen grains is due to the resistant properties of the
outer coat of the grains, when the pollen is deposited out of reach of the oxydizing
action of the air. This condition is particularly fulfilled in peat bogs, with the
result that pollen grains are found most abundantly in peats, browncoals and
lignites, and coals.
In the half a century following G6ppert's first observation of fossil pollen
grains, such microfossils were also studied by, among others, the famous C. G.
Ehrenberg, an early pioneer of micropalaeontology, and by the Swiss workers
F. E. Geinitz and J. Friih.
A very notable early contribution to the study of ancient microfloras is
the two-volume work of Paulus REINSCH, Micro-Palaeophytologia Formationis
Carboniferae, published in 1884. Most of the illustrations in this work were made
in black crayon and are so clear that they are only bettered by the best modern
photographs (Fig.l).
When TRXBOM (1888) encountered pollen grains of pine and spruce in a
Quaternary lake deposit in Sweden, he considered these to be useful index fossils.
Further important contributions were made by C. A. Weber, a German peat
stratigrapher, and his school (WEBER, 1893, and later). It is in his work that a
quantitative presentation of pollen-analytical data is found for the first time,
and already in relative figures.
U. Steusloff, while studying certain terraces, peats and lake deposits in
northwestern Mecklenburg, Germany, in the first few years of this century,
took a particular interest in microfossils occurring in certain deposits of lake
lime. He found that staining with Magdala red made pollen grains stand out
in a brilliant red colour. He studied samples from seven strata of alternating
light and dark lake marl, and counted from each sample between five hundred
and seven hundred pollen grains. Steusloff believed that the banding was of
seasonal origin with spring layers, poorer in lime, alternating with autumn layers,
richer in lime; but the pollen content of both was nearly the same, despite the
fact that he noted that the pollen grains shed during spring and early summer are
entirely different from those shed later. Steusloff, therefore, suggested that pollen
grains deposited in lake waters may float in the water some time before sinking
down to the bottom (see further ERDTMAN, 1943, pp.6-7).
Fig. I. Plate XXII from Paulus Reinsch, Micro-Palaeophytologia Formationis Carboniferae (1884).
H A L F A C E N T U R Y OF MODERN P A L Y N O L O G Y 281
el (~m O-
7
~,tOm Jt
dlw~ q ~
282 A.A. MANTEN
In Finland, at about the same time, H. Lindberg found that the percentages
of certain pollen forms can differ in successive layers.
Another important investigator of fossil pollen was the Swede Gustav
Lagerheim. Although he himself did not publish anything on pollen analysis,
his contribution to this branch of science should not be underestimated. Results
of analyses made by Lagerheim were published by WITTE (1905), HOLST (1909),
SAMUELSSON (1910), SERNANDER (1911), VON POST (1918) and the Swedish Geo-
logical Survey. It was through Lagerheim that Von Post learnt to see the strati-
graphical value of plant microfossils.
Subdivision of the Postglacial and Holocene
For a correct understanding of the historical development of pollen analysis,

it is necessary to say a few words here about attempts to establish a chronological
subdivision of the period between the end of the last glaciation and the present
time.
As early as 1841, the Dane Steenstrup made a peat study that led him to
distinguish a stratigraphical succession of a Populus tremula zone, a Pinus zone,
a Quercus zone and an Alnus zone. He ascribed this zonation to a gradual im-
provement of the climate which has taken place since the last glaciation (SrEEN-
STRUP, 1842).
In 1876, the Norwegian BLYI'T presented a theory about climate in his
country since the retreat of the ice cover. Based upon the penetration in Norway
of the present flora, he distinguished a few alternating warmer and cooler, dryer
and wetter periods.
His theory was further extended by the Swede SERNANDER (1908, 1910),
who made it more generally applicable. Since then the successive periods have
been known for a long time as the periods of Blytt-Sernander.
Pollen in honey
A fourth line of pollen studies began in 1895, when the German PFISTER showed
that the geographical and botanical origin of honey can be determined by means
of the pollen grains which are found in the honey. When a bee visits a flower, it
also comes in contact with the anthers. As a result, some ripe pollen usually
drops on the nectar, and is taken together with this nectar to the honey combs
in the hive, from where the apiarist collects the honey to market it.
Initially Pfister's discovery stood more or less by itself. It was more than
a decade before his work was followed up by studies of other researchers. These
were YOUNG (1908) and FEHLMANN (1911), who worked on American and Swiss
honey, respectively. After their publications, a hiatus again occurred in melitto-
logical pollen studies, which lasted until the early nineteen-thirties.
Earth-Sci., Rev., 2 (1966) 277-316

LENNART VON POST
One of the scientists, who decidedly influenced the life of Lennart yon Post,
was the colourful and stimulating Rutger Sernander. It was he, in 1902, who gave
the geology student Von Post the first scientific assignment: to prepare a report
about the historical development of the M~istermyr, one of the main marshes
of the Baltic island of Gotland, for which a reclamation plan had been designed.
Afterwards, Von Post kept his sincere interest in peat studies. The thesis
for his master degree which was presented at the University of Uppsala, in 1907,
dealt with peat swamps in Norrland. In 1908, he joined the staff of the Swedish
Geological Survey and began to work on swamps in the province of N~irke (west
of Stockholm). In this investigation he began to use pollen grains as stratigraphical
indicators, and he introduced a "spruce-pollen boundary" as a basis for local
correlations. This boundary, however, did not help him in later work on peat
swamps in Scania, and finally the only way out appeared to study the strati-
graphical distribution of all arboreal pollen. The results were so stimulating
that Von Post decided also to revise his material from N~irke, identifying and
counting all arboreal pollen grains found therein 1.
Von Post arranged his results in graphs, extending the depths of his samples
along the vertical axis and the percentages of the various pollen species found
in these samples along the horizontal axis. (In a later stage, the plotted points
were indicated by means of standard signs, each sign always indicating the same
species.) Lines were drawn to connect the successive points. The course of each
line thus was a reflection of variations in abundance of a particular tree in the
course of time of peat formation. Such a graph was called a pollen diagram.
The first demonstration of pollen diagrams to a large scientific public took
place in a lecture at the Sixteenth Scandinavian Meeting of Natural Scientists,
in Christiania (now Oslo), in 1916.
It was not entirely coincidental that pollen analysis was developed in Scan-
dinavia, in an area which had been glaciated during the Pleistocene and which did
not exhibit topographical height differences of much importance. In that area,
the forests are composed of only a limited number of tree species, each of which
produces much pollen. The smooth topography meant that alterations in the
composition of the forests could be recorded almost unhindered in the fossil-
pollen spectra. In a mountain area, and with a much richer and more varied
vegetation cover, the picture presented by fossil pollen is much more complicated.
The fact that, later, successful studies have also been made of such areas is another
matter entirely, as then the principles of the method were known.
Von Post's method of pollen analysis was not received immediately with
general appreciation. It seemed doubtful to several people that pollen grains
1 F o r a more detailed review of the development of the peat studies of Von Post, see MANTEN
(1967a).
Earth-Sci. Rev., 2 (1966) 277-316

284 A.A. MANTEN
which had often been transported over long distances, could be used as a trust-
worthy basis for establishing a stratigraphy of the bogs in wihch they finally had
been deposited and preserved.
It was mainly for other reasons that Von Post was asked to continue his
peat studies. World War I was going on and Sweden was facing a difficult fuel-
supply situation, since it had neither coal nor petroleum of its own. This focused
attention on the peat resources of the country and Von Post had to make a survey
of the peat bogs in southern Sweden, as a basis for calculations of the amount and
value of the nation's peat resources (VoN Posx and GAVELIN, 1917). The results of
pollen-analytical studies carried out in connection with this inventory and later
investigations, as well as early studies by other scientists helped significantly
to overcome much of the initial mistrust of the reliability of the pollen-analytical
method.
THE RISE OF MODERN PALYNOLOGY
Quaternary pollen analysis
A stimulating discovery in the early days of pollen analysis was that the various
periods of the Blytt-Sernander chronology of the Postglacial and Holocene
showed up very well in pollen diagrams. Thus, the following periods could be
distinguished, starting from the retreat of the last glaciation: a Subarctic period,
a Late Glacial period, a Preboreal period, a Boreal period, an Atlantic period,
a Subboreal period, a Subatlantic period, and the present time, which is often
thought to be part of the Subatlantic period still. It was gradually realized,however,
that no great value could be attached to the climatological importance which
Blytt and Sernander awarded to their periods.
Following the example of Blytt and Sernander, several other subdivisions
of the Holocene have been published, based upon standard pollen diagrams of
a certain area. A well-known one is that by RtJDOLPH (1931), who subdivided the
forest history in central Europe into, successively: (1) Betula-Pinus period, (2)
Corylus period, (3) Quercetum mixtum period, (4) Fagus period. In more re-
stricted areas local variations may occur such as the intercalation of a Picea period.
Other subdivisions of the Holocene are arranged with numbered zones, such as
those proposed by JESSEN(1934, 1938), OVERBECKand SCHNEIDER(1938), SCHUT-
RUMPE (1943), and FIRBAS (1949). As far as the subdivision of the Early Holocene
is concerned, these later chronological subdivisions present no essential improve-
ment over the classical chronology. However, the subdivision of the Late Holocene
is better in these later chronologies than in that of Blytt and Sernander.
Von Post lived long enough to see the results of pollen-statistical studies
extend knowledge about Quaternary plant geography in a revolutionary way
and the method become an indispensable stratigraphical aid. On several occasions,
Earth-Si. Rev., 2 (1966) 277-316

he presented reviews about Quaternary pollen analysis (VON POST, 1924, 1930).
In the latter contribution, he demonstrated how pollen diagrams can be used
to correlate deposits from all over Europe.
In the first two decades, pollen analysts mainly followed the classical lines
set out by Von Post, gradually expanding the area of investigation, especially
in those parts of the world where the problems were essentially similar to those
in Scandinavia. The investigators were principally engaged in solving primary
problems of forest history, shore-line movements, etc.
An important improvement of the method was the suggestion by FIRBAS
(1935) to pay attention also to the values of the percentages for the herbaceous
pollen grains, particularly that of the Cyperaceae, Ericales and Poaceae (Gra-
mineae). In this way it became possible to recognise, palynologically, forest-less
periods with tundras and periods with a vegetation with scattered small occur-
rences of trees and bushes in a forest-less territory (for the latter kind of vege-
tation Iversen later introduced the name of park tundra), and also to study
the history of the moorlands.
Firbas initially had the idea that during the last, the Wiirm Glaciation,
the northern ice cover was surrounded by a tundra, around this a park tundra,
next a subarctic forest with birches and firs, and finally around this the forests
with broad-leaved trees. All these kinds of vegetation would thus have existed
simultaneously, but at different places, depending on the geographical distance
to the ice cap. It was also assumed that with the retreat of the land ice, in one
area these kinds of vegetation succeeded each other in time.
It was shown in Denmark that this idea of a gradual improvement in climate,
reflected by the succession in time of the named vegetations, was incorrect. Pollen-
analytical studies made it clear that during the kate Glacial a warmer stage inter-
rupted a period which was on the average still rather cold. This interruption was
called the Allerod oscillation. Continued research showed that this oscillation
had also occurred elsewhere in Europe, thereby influencing plant growth. As
a result, JESSEN (1935) published a new subdivision of the Late Glacial in which
the Allerod oscillation was officially included.
It should be said here that even before this, in Denmark the idea had been
launched that the improvement of climate since the retreat of the land ice had
not been as gradual as one might be inclined to think. HARTZ and MILTHE~S
(1901) had found in Late Glacial deposits an alternation in the nature of sedimen-
tation which they believed must have been caused by a climatic fluctuation.
Probably because they published their conclusion in Danish and because it
came from another field of study, palynologists did not pay much attention to
the article of Hartz and Milthers until they found the oscillation themselves
in the pollen diagrams. Then, however, they honoured the pioneering work of
Hartz and Milthers by naming the oscillation after the site from which the type
deposits were first described.
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286 A . A . MANTEN
Pollen analyses and archaeology
In addition to making relative datings of peat horizons, pollen analysis can also be
used to determine the age of prehistoric objects found in such deposits. This
made the method also of great value to archaeology. On several occasions Von
Post helped to make relative-age determinations of archaeological findings by
means of pollen analysis. The most well-known of these is the dating of a bronze-
age mantle found at Gerumsberg, in V~isterg6tland (VON POST, 1925).
In 1937, FIRBAS, by recognizing the pollen of cultivated cereals, showed
that pollen analysis could be applied to the study of the prehistorical development
of agriculture. A year later, FROMM, through a study of material collected by
Lid6n, succeeded in correlating the results to known geochronology.
Palynology and coal geology
New impulses were brought into palaeopalynology when it was shown that it
could be used for the identification and correlation of coal seams and coal-
bearing rocks as well. In 1920, the American geologist TmESSEN assumed that
spores from Pennsylvanian coals might be used for stratigraphical correlation.
Proceeding with this idea through a study of thin sections of the "Thick Free-
port" coal, THIESSEN and VOORHEES (1922) found that, where the spore content
is concerned, "there is a variation in the composition of the bed vertically, but
none horizontally". They also noted that several of the spores found in the
Freeport coal bed were different from the spores which they had observed in
other coals. Identification of various coal deposits by means of characteristic
spores and plant structures was, consequently, elaborated in later papers (THIESSEN
and STAUD, 1923; TmESSEN and WILSON, 1924).
Once it had been found possible to correlate coal beds in this way, similar
investigations were also carried out in other countries, and the technique was
especially expanded and refined in central Europe. LAN6E (1928) examined and
plotted the stratigraphical distribution of Carboniferous megaspores, small
spores, and fungal remains found in German and Polish coals, using thin sections
and isolated-spore preparations. He obtained isolated spores by crushing the
coal to about 1.5 mm, mixing it with a mixture of CC14 and xylol with a s.gr. of
1.5; centrifugation of this mixture resulted in the float containing the lighter
spores which were freed from the coal when it was crushed. Another pioneer
was ZERNDT, who in the nineteen-thirties investigated Carboniferous megaspores
from Polish coals using the maceration method.
It was also particularly in central Europe that the application of palynology
to the study of browncoal and lignite deposits was developed. Closely connected
with this field is the name of R. Potoni&
Probably as a result of the fact that coal, browncoal and lignite represent
Earth-Sci. Rev., 2 (1966) 277-316

ancient peats, and peat was the preferred deposit of pollen analysts in the first
few decades after Von Post, little attention was paid to the spores-and-pollen
content of the sediments underlying or overlying the coal. Once they were also
included in the studies, mainly after World War II, these, particularly the clays,
were often found to contain a richer and more varied microflora than the coal
itself.
The nomenclature problem in Tertiary palynology
HECK (1927) seems to have been one of the first to undertake a study of Tertiary
pollen in this century (el. KmCHHEIMER, 1940). Several others soon followed.
As we shall see in later pages, it was the application of palynology in petroleum
exploration that has given Tertiary palynology its greatest impetus. This began
slowly in the nineteen-thirties, but the real "boom" only came after World War II.
Whereas Quaternary palynologists could identify all pollen grains which
they found in terms of present-day taxa and Palaeophytic palynologists had to
work throughout with organ taxa, Tertiary palynologists had to deal with an inter-
mediate situation. As the sediments under investigation are progressively older,
less and less of the pollen can be correlated, with certainty or with approximation,
with modern botanical genera.
In this situation, a series of publications by Potoni6 and co-workers has
greatly influenced Tertiary palynology (C. A. BROWN, 1957). These show many
developmental changes in concepts as well as in nomenclature.
In 1931, POTONII~started by placing fossil spores and pollen grains in the
form genera Sporites and Pollenites. Since the chances of finding fossil pollen
grains or spores of a plant are at least as great as that of finding any other fossilized
organ, it soon became evident, however, that two form genera were definitely
insufficient and that a much higher number of form genera was required, even
though pollen or spores of many species and sometimes also higher taxa are not
sufficiently distinct to be kept apart. Consequently, Potoni6 dropped the two-
form-genera system after only a few years. Instead he then began to classify
fossil pollen grains and spores within the framework of a botanical classification
(PoTONII~, 1934). He initiated the system of adding "pollenites" or "sporites"
as a hyphenated suffix to the generic name, as in Alni-pollenites verus. However,
this system was not satisfactory in all respects, for in many cases he could not
give a positive generic identification. He indicated the uncertainty with a question
mark, as in Coryli?-pollenites coryphaeus.
WODEHOUSE (1933) suggested contracting the suffix "pollenites" to "pites"
and to add this to the specific designation if the genus of the pollen species is
known with the normal degree of accuracy, otherwise to its generic designation
(e.g., Pinus strobipites, Ericipites longisulcatus).
THIERGART (1940), being faced with the same problem as Potoni6, simply
Earth-Sei. Rev., 2 (1966) 277-316

288 A.A. MANTEN
used existing generic names for the fossil spores and pollen grains which he
felt able to correlate positively to present-day genera and used the form genera
Sporites and Pollenites for the others.
Still not satisfied with the attempts then made in the classification and
nomenclature of fossil spores and pollen grains, Potoni6 proposed in 1950, to-
gether with Thomson and Thiergart (POTONI~ et al., 1950), the system of adding
the suffixes "-oidites" to generic names and "-oides" to specific names (e.g.,
Quercoidites, Fagus silvaticoides), in this way showing to what taxonomic rank
they felt reasonably certain about the identification of fossil pollen. This system
has been designated the "half-natural system". In 1951 and 1959, Potoni6 again
altered the nomenclature by altering names such as Quercoidites to Quercoi-
pollenites; these categories he apparently considered as organ genera.
The changing of the system of classification from a form genus, Pollenites,
to one with suggested botanical relationships, and then from a half-natural sys-
tem to one using organ genera, caused much confusion. THOMSON and PFLUG
(1953) recognized this, so they established a system of nomenclature based on
the morphology of pollen, which is a system of form genera. But they did not
preserve their type specimens. Part of their system is based on the system of
classifying modern pollen into artificial sections, as proposed by IVERSEN and
TROi~Ls-SMITH (1950; see also FAEGRI and IVERSEN, 1950).
INGWERSEN (1954) on the other hand, used present-day botanical genera
to classify many of the pollen grains which he found in Danish lignites.
All these nomenclature systems together are still in use at the present day.
The haphazard application of artificial names, "half-natural" names and
botanical names to fossil pollen is deplored by several palynologists. For the time
being the best practical solution seems to be to use one artificial system for all
Tertiary and older pollen, so that each kind of fossil pollen has only one name
(MANTEN, 1958, 1965). However, such a system conflicts in some respects with
the existing International Rules of Botanical Nomenclature. Under these rules
we can, for instance, have no organ taxon which is synonymous with a taxonomic
unit. A fossil pollen grain referable to, e.g., Alnus, is an Alnus pollen grain and
not Alnipollenites; should the identification not be quite certain, the pollen grain
may be called a cf. Alnus pollen grain.
Palynology and the petroleum industry
Most petroleum companies initially showed great reluctance to make use of

palynology in their geological-prospecting work. Fossil plants had up to then
never contributed much to the resolution of marine and brackish-water strati-
graphical problems, and, as a result of this, more or less, petroleum geologists
had seldom had any palaeobotanical training.
A change in this situation began very gradually in the middle of the nine-
Earth-Sci. Rev., 2 (1966) 277-316

teen-thirties. It did not become very apparent, however, until after World War II,
and particularly in the nineteen-fifties.
The first attempts to find out whether palynology could be useful to the
geological activities of petroleum companies seem to have started in 1934, in
the U.S.A. In some areas, petroleum geologists were hampered in their work
on pre-Quaternary, particularly Tertiary, series of strata, because existing palaeon-
tological methods of determining age and correlation could not be used and also
lithological methods could not be employed with sufficient results. Thus, they
began to look for some other way to obtain the desired information. The suggestion
was made that perhaps pollen analysis could solve the problems. The question
was tackled mainly by expert consultant advice and some preliminary investi-
gations by palaeontologists in the companies' own service. Thus, in 1938 and
subsequent years, the Royal Dutch/Shell Group invited such specialists as Potoni6,
Florschfitz, Stiitzer and Bode to undertake a study of Tertiary material from
Mexico, Venezuela, Trinidad, the Far East and Columbia, on a consultant basis,
and also had their own palaeontologist T. F. Grimsdale do some pioneering
palynological work. The main problem was that knowledge of Tertiary pollen
had at that time not yet reached the same stage as that of Quaternary pollen.
Nevertheless, the first results were promising enough to lead the Royal Dutch/
Shell Group to decide to establish a palynological section within its stratigraphical
department. Because of World War II, however, this could not take place until
1946.
Botanical microplankton and chitinozoans
The same treatment as is used in the preparation of samples of fossil spores and
pollen grains, can also be used for other groups of microfossils, such as micro-
plankton of botanical affinity, chitinozoans and other types of algal and protistan
entities. Gradually, study of these objects also became part of palynology. The
procedure used is then taken as the boundary with older branches of micro-
palaeontology, which make use of the techniques of alkaline washing of disinte-
grated rock, such as in the study of Foraminifera or Ostracoda, or of sectioning
of hard rocks.
Microplankton
Fossil microplankton organisms of botanical affinity were first described by
C. G. Ehrenberg, the father of micropalaeontology, in the eighteen-thirties (cf.
EHRENBERG, 1836, 1854). During the half century following his pioneering studies,
such microfossils were the subject of only rather casual mention in a limited
number of scientific publications. In the early decades of the twentieth century,
fossil phytoplankton assemblages were recorded from several countries.
This situation only really effectively changed in the first half of the nineteen-
Earth-Sei. Rev., 2 (1966) 277-316

290 A.A. MANTEN
thirties, when several European palaeontologists, among whom O. Wetzel, A. Ei-

senack, G. Deflandre and M. Lejeune-Carpentier, began an important series
of studies of these botanical microfossils, which has continued until the present
time.
For all spherical spinose remains that were found among the fossil micro-
plankton, the name hystrichospheres was introduced. This general term, literally
meaning "spiny balls", was derived from two form genera, Hystrichosphaera
O. WETZEL and Hystrichosphaeridium DEFLANDRE, and was widely used in
scientific literature from the nineteen-thirties until the fifties.
Chitinozoans
The discovery of the group of microfossils known as Chitinozoa is linked to
the studies of Eisenack, which aimed at tracing glacial erratics found in northern
Germany back to their sources. In this connection, he investigated the acid-
resistant microfossils found in the erratics and in Silurian deposits of the Baltic
a r e a (EISENACK, 1931, 1932, 1934, 1938). Among these fossils, he noted a group
of small (ca. 0.15-1.5 mm), thin-walled, flask-shaped vesicles, which had not been
described earlier in the literature. Because of the resemblance of their wall to
chitin, Eisenack named this group of fossils Chitinozoa. He also developed a
systematic nomenclature, which remained in use essentially unaltered until in the
nineteen-sixties. His classification was based on overall morphographical char-
acteristics of silhouette and ornamentation, and ratios of measurements. His
diagnoses did not contain information on internal structures and wall construction,
for his material consisted mainly of opaque specimens. Until 1940, he recognized
some twelve different genera of Chitinozoa. The systematic position of the order
Chitinozoa is uncertain, but it is generally assumed that they represent remains of
animals.
Pollen studies in the service of medicine
It has already been mentioned that Scheppegrell revived the "gravity-slide"

method of systematic pollen trapping, in 1916, a very important year in the
history of palynology. This method has been applied in the U.S.A. probably
every year since and in almost every state of the country. Similar studies followed
in other countries, particularly in Europe, South America, New Zealand and
Australia, and South Africa. The paucity of observations from tropical areas is
due presumably not to the non-occurrence of hay fever and hay asthma, but
only to their relative unimportance, real or supposed.
A more ideal method than that of using a horizontal surface coated
with an adhesive, as in the gravity-slide method, would be to determine at short
intervals the load in pollen grains per unit volume of air and several forms of
Earth-Sci. Rev., 2 (1966) 277-316

apparatus have been devised for this purpose. Most aerobiologists, however,
went on trapping pollen on horizontal slides.
The aerobiological studies have given much more insight into which pollen
cause inhalent allergies. Any pollen, qualifying as a probable cause of epidemic
hay fever and hay asthma, must answer certain requirements: (1) it must be anti-
genic; (2) is must be produced in large quantities per plant by a species which
is widely and abundantly distributed and is anemophilous in habit; (3) the pollen
of such a species must be sufficiently buoyant (in practice its grains must be small
enough) to become easily airborne. To these requirements must be added that
for any type of pollen to be regarded as the causitive agent in a particular case
of hay fever or hay asthma, its period of occurrence in the air must coincide
with or include that in which the patient's symptoms are experienced, or at least
are exacerbated ("time postulate") (HYDE, 1954).
The "palynological" problem presented by hay fever was to determine what
kinds of pollen answering the above requirements were likely to be encountered
in the air at a given place in the course of the year, during what period and in what
concentration(s), and how such incidence varies from place to place.
Treatment of pollen allergies

As mentioned before, the oldest treatment of hay fever is bringing the patient
to a place where the air is free of the causative pollen. A closely related treatment
is that of modification of the patient's daily time table, taking into account that
part of the 24-h cycle when the pollen causing the hay fever is most abundant
in the air. This method gained in importance with increasing aeropalynological
knowledge.
Another approach is that of desensitization of the patient to the causative
pollen with pollen extracts. After he had discovered the nature of hay fever,
Noon was the first to apply preseasonal treatment to his patients. WALKER(1921)
noted that in cases where such treatment led to little or no improvement, pollen-
extract injections may be continued, but in smaller amounts (co-seasonal therapy).
WALKER(1920) was also the first to suggest that if pollen cases were treated long
and consistently, many patients would possibly be entirely free from symptoms
for years, if not throughout life. The first two comprehensive papers which
reported good results of such perennial treatment were by A. BROWN (1927)
and by VANDZRVEZR et al. (1927). In the next few years others reported success
as well, and therefore such treatment also became an accepted procedure.
Melittopalynology
The study of the pollen found in honey began to flourish again in the early nine-
teen-thirties. The microscopic examination of honey was then developed mainly
in Germany, by Armbruster and co-workers (ARMSRUSTER and OEN~KE, 1929;
Earth-Sei. Rev., 2 (1966) 277-316

292 A.A. MANTEN
AR~BRUSTER and JACOBS, 1934~1935), GRIEBEL (1930--1931), and especially by

ZAYDER (1932, 1935, 1937, 1941, 1949, 1951), who through his enormous amount
of work laid down the foundations of the microscopic determination of the origin
of honey. During the following years, the microscopic analysis of honey was
also developed in various other countries of Europe; regional types of honey
and the pollen combinations which characterize them were described. Critical
studies also showed the possibilities and limits of the method.
Pollen morphology
After the great advances in pollen morphology made in the nineteenth

century, it was only in the nineteen-thirties when further really significant contri-
butions to this field appeared.
In his studies of the browncoal deposits in the Lower Rhine area, Potoni6
found many pollen grains which were unknown in literature and, therefore,
needed to be described in detail. However, there was such a variety in character-
istics that he felt it necessary to give first an explanation of his terminology
(POTONII~, 1934). This publication reveals a great insight into the construction
of the pollen-grain wall. With the aid of this terminology, Potoni6 described the
fossil pollen grains and his system is still used by several palaeopalynologists.
A number of terms used by him were adopted from Fritzsche, others from Wode-
house, who had already started to publish information about pollen grains in
1928.
In contrast to Potoni6, Wodehouse mainly studied recent pollen and even
mainly fresh grains. His studies were made primarily to support allergy research.
For North American physicians, it was most important to find out as soon as
possible what kind of pollen caused hay fever in a certain area with a certain
patient. This was only possible when good descriptions of the pollen grains were
available. The work of WODEHOUSEculminated in a book which appeared in 1935.
For his descriptions, he also developed a system of terms, some of which were
again adopted from Fritzsche, but many others were new. In addition to the
description of pollen grains, Wodehouse also paid much attention to the devel-
opment and arrangement of apertures, and to the coherence of the pollen tetrads.
Surface structures could not be studied well, as they were masked by the contents
of the pollen grains (protoplasm, intine). He did not apply the method developed
by the brothers G. and H. Erdtman of treating pollen grains with strong acids,
which dissolved the pollen content.
G. Erdtman approached pollen morphology from quite a different angle.
He was a student of Lennart von Post, and in the nineteen-twenties was mainly
engaged in pollen-analytical work. He realized that the study of pollen grains
from a morphological point of view had lagged behind the results of pollen analysis,
which in fact was based on pollen morphology. "It is, however, true that many
Earth-Sci. Rev., 2 (1966.) 277-316

of the specialists most interested in the results of pollen analysis have not had a
direct professional interest in pollen morphology or the fundamental theory of
pollen analysis" (ERDTMAN, 1943). For this reason, in his introductory textbook,
he devoted much attention to pollen morphology. The system of terms which
Erdtman used was partly based on the terminologies of Fritzsche and Wodehouse,
but many new terms were also introduced. For the first time all pollen grains were
given consistent chemical treatment according to the acetolysis method of the
brothers Erdtman. The pollen-grain wall, in particular, could thus be studied
better than ever before. Recent pollen treated in this way closely resembles fossil
pollen; much more so than fresh material.
First books on palynology
The first book on palynology appeared in 1935. It was WODEr~OUSE'Swell-known

Pollen Grains. Their Structure, Identification and Significance in Science and
Medicine. In 1942, the Lehrbuch der Pollenanalyse, by K. BERTSCH, appeared,
which treated pollen analysis in general, but also presented a guide to the fossil
pollen grains and spores and leaf remains, as well as some of the more common
animal microfossils. In 1943, ERDTMAN'SAn Introduction to Pollen Analysis appeared
and had a good reception as a most needed treatise of the subject in the English
language.
A DISCUSSION ABOUT THE RIGHT NAME
Because World War II caused the suspension of many scientific meetings and
handicapped travelling, P. B. Sears felt the need for a medium to exchange
information among U.S. workers in pollen analysis. On May 5, 1943, in col-
laboration with some others (L. R. Wilson, J. E. Potzger), he started the publi-
cation of a mimeographed Pollen Analysis Circular. This was the first periodical
entirely devoted to pollen analysis.
In the sixth issue of this Circular, ANTEVS (1944) queried whether "pollen
analysis" is the proper name for the study of pollen, and its applications. The
word, he said, was used in Sweden from the beginning to signify the identification
and percentage determination of the pollen grains of the principal forest trees
in peat bogs and lake beds. However, its inadequacy soon became obvious.
Erdtman spoke about "pollen statistics", and, after 1932, about "pollen statistics
and related topics". Even the combination "statistical pollen analysis" was rejected
by Antevs, since it "refers only to the method of getting certain data which in
itself has little purpose and which does not apply to or cover all the branches
of the pollen studies, much less the application of the direct results to climatic
conclusions, etc. It is the knowledge gained from the pollen studies, be those
Earth-Sci. Rev., 2 (1966) 277-316

294 A. A. MANTEN
statistical or morphological, or be they concerned with pollen-induced diseases

as hay fever, etc., that has purpose and significance."
After concluding that a name like "pollinilogy" would not be very good
either, Antevs ended with suggesting "pollen science", a suggestion which,
according to a postscript, appealed so much to the Editor of the Circular, that
he suggested renaming the periodical the "Pollen Science Circular".
In the next issue, WILSON (1944) applauded the suggested new name of
the publication, but pointed out that those engaged in Palaeozoic and Mesozoic
studies work with spores as much as with pollen and other microfossils. He pro-
posed that "we could use the word 'pollen' in a very liberal sense and understand
that spore studies were to be included in the Circular" (and, as he no doubt
meant to say, in the science).
In reaction to this, HYDE and WILLIAMS (1944) wrote that in such a case,
some word carrying a wider connotation than pollen seemed to be called for.
They suggested palynology, derived from the Greek r~0tk,3wo (palyno), which
means to strew or sprinkle, and is suggestive of n~)~'q (pal6), fine meal, which is
cognate with the Latin pollen, flour, dust. Palynology, they defined, is the study
of pollen and other spores and their dispersal, and applications thereof. They
ventured "to hope that the sequence of consonants p-l-n (suggesting pollen,
but with a difference) and the general euphony of the new word may commend it
to our fellow workers in this field", a hope which, as history proves, has been
richly honoured.
The suggestion of "micro-palaeobotany", made in the same issue of the
Circular (BRYAN, 1944), has hardly been discussed.
o
-; 45OO
4ooQ
]3
350C
3_
300(
~5
250(
>~ 2 0 0 (
150(
E
1000
500 ~
,
1935 1 9I4 0 I
1945 19 I5 0 19155 I
1960
Fig.2. Rate at which new species and emendations have been added to thepalynological literature
in the years 1931-1956. (After KREMPet al., 1957.)
Earth-Sci. Rev., 2 (1966) 277-316

In the ninth issue, which appeared on January 15, 1945, Sears, Editor of
the Circular, professed to be "intrigued with the suggestion of Professors Hyde
and Williams that the term palynology be used to designate the whole science
which deals with strewn or scattered organic particles". Recipients of the periodical
seemed to be more hesitant in accepting the new name and the majority suggested
calling it Pollen and Spore Circular, a name which it bore from the ninth to the
eighteenth, and last issue.
The discussion ended in the tenth issue, when HYDE (1945) wrote that
Dr. Williams and he did not intend the word palynology to have so wide an
application as Sears had given it. He admitted that he could not object on logical
grounds to the inclusion of fungus and bacterial spores, but would certainly
not wish to include, e.g., virus particles or dead organic dusts.
THE MATURITY OF PALYNOLOGY
Palynology and the petroleum industry
There is no doubt that it is the successful utilization of plant microfossils in

stratigraphical work of petroleum geology that has given modern palynology
its greatest impetus. In 1946, the Royal Dutch/Shell Group added a palynological
section to its stratigraphical department, and a year later the Group established
its first operational palynological laboratory, in Maracaibo, Venezuela. In the
period 1945-1947, also two other major companies--Creole and Carter (Jersey)
--worked independently to solve stratigraphical problems by means of palynology,
and were successful in establishing correlations.
Thus, the first few years after World War II opened a most important era
in petroleum palynology. During the first half of this decade, the growth of paly-
nology in the petroleum industry was still slow. It was characterized by the be-
ginning of recruitment and training of palynologists in the full services of petro-
leum companies, and the attack on pre-Quaternary deposits. This led to the break-
through in the palyno-stratigraphy of the Tertiary and the subsequent mushrooming
of petroleum palynology. Vast amounts of money were spent during the nineteen-
fifties in establishing palynological laboratories to assist petroleum companies
all over the world. By the end of that period, almost every major company and
many independents had such laboratories.
Until about 1956, the main activities of petroleum palynologists concerned
the Tertiary. From 1956 onwards, however, growing interest was shown in
Mesozoic and Palaeozoic sediments, but it was only in very recent years that
stratigraphical work on pre-Cenozoic deposits superseded petroleum-palynolog-
ical work on the Tertiary in importance. The main stimulating factors were
Earth-Sci. Rev., 2 (1966) 277-316

296 A.A. MANTEN
increasing applications of palynology in petroleum-exploration work in the

Middle East and in European off-shore areas.
The prolific growth of palynology in the years 1951-1959 is partly re-
flected by the increased output in scientific palynological literature. Whereas
during the first 35 years of modern palynology (1916-1950) about two thousand
scientific publications appeared, dealing with this branch of science (SELL1N6,
1951) an almost equal number of publications was produced by the end of the
nineteen-fifties in the space of less than three years. If, in addition, we realize
that to an increasing degree petroleum companies were collecting unpublished
palynological information, it may be assumed that in the years 1950-1959 palyno-
logical activity expanded with a factor probably of about 10. The increase is
reflected in, among other things, the rate at which new species and emendations
have been added to the literature (Fig.2).
Since 1959, this rapid development of palynology has slowed down to a
more normal yearly increase. Each year, roughly about 5-10~o more publications
are produced than in the previous year. Again taking into account unpublished
work, it may be estimated that at present the total palynological activity doubles
in about 5-7 years.
A critarehs, dino.flagellates and chitinozoans
It has already been pointed out that in the decade preceding World War II fossil
botanical microplankton, chitinozoans and similar entities began also to be used
in palynological studies, in addition to the expanding amounts of work on spores
and pollen grains. For the entire group of palynological study objects, Tschudy
introduced the collective name of palynomorphs.
Attempts to clarify the systematic nomenclature of fossil microplankton,
that had grown into a confused mass of uncorrelated data, revealed that it is
possible to recognize fossil chorate cysts of dinoflagellates and to separate them
from other microplanktonic bodies incertae sedis. These cysts had been de-
scribed at various times by different names, such as "Xanthidia", "Spiriferites",
"Palinospheres", "Ova hispida", "Hystrichospheres" and "Hystrix". The first
to realize that these were dinoflagellate cysts was REINSCH (1905). Four decades
later, DEFLANDRE(1947a, b) also gave evidence in favour of this interpretation;
he established beyond any doubt their truly planktonic character. The matter
was definitely solved by EVITT, in papers published between 1961 and 1963. It is
very ironic that both the genera Hystriehosphaera and Hystrichosphaeridium,
from which the group name of the hystrichosphaerids was derived, happened
to be based on fossil dinoflagellates. A short historical review of the study of
fossil dinoflagellates is given by SARJEANT (1967) and a bibliography of papers
published on this subject up to 1963 is given by DOWNm and SARJEANT(1964).
In order to eliminate future confusion between fossil dinoflagellates, hvs-
Earth-Sci. Rev., 2 (1966) 277-316

trichospherous or otherwise) and other microplanktonic fossils (probably mostly

of algal affinity and often with spinose ornamentation or structure), EVITT, in
1963, proposed calling the latter group Acritarcha--derived from two Greek
roots and meaning "of unknown origin". This term has found ready acceptance,
even though some authors used the hybrid, but more descriptive "acritarchous
hystrichospheres" rather than "acritarchs". The natural affinities of the several
groups within the Acritarcha are largely unknown. This is partly because few
studies of Recent marine microplankton had reported the presence of bodies
resembling the abundant fossil acritarchs. In fact, some palynologists were in-
clined to believe that acritarchs had become extinct during the Late Tertiary.
This, however, has changed during the last few years. With the new stimulus
given to planktology by the need to modernize and intensify the fishing industry,
several isolated reports of the existence of living Acritarcha have been published.
There is, thus, hope that the natural affinity of this "acritarchous" group of
fossils that occur in deposits from the Precambrian onwards and are abundant
since the earliest Palaeozoic, will still be learnt.
The reason why interest in various palynomorphs other than spores and
pollen grains has increased rapidly especially during the last decade is roughly
the same as that leading to the sudden sharp growth of pollen-and-spore analysis
earlier, viz. because their potential use in stratigraphic palaeontology had become
widely appreciated.
As has been mentioned before, acritarchs can be used from Cambrian
deposits onwards. Although dinoflagellates are known from the Palaeozoic and
perhaps were already in existence even in the Precambrian, they are mainly
important tools for the correlation and dating of deposits from the Middle and
Upper Jurassic, Cretaceous and Tertiary. Chitinozoa are of great and increasing
stratigraphical value in deposits from Ordovician to Devonian in age.
Quaternary palynology
In 1946, IVERSEN published a new method of presenting pollen-statistical data.

In his calculations, he treated the arboreal pollen on equal terms with the pollen of
anemophilous herbs and Ericales. The total of the pollen grains counted for
these groups together constitutes the basis for the calculation of percentages.
Pollen diagrams constructed from this information, with arboreal pollen plotted
from left to right and the herbaceous and Ericales pollen from right to left,
immediately show variations in forest density. For statistical reasons, this method
required the counting of about 500 pollen grains per sample.
lversen also introduced the concept of arboreal-pollen frequency, that
is the average number of arboreal pollen grains per slide. In this way, too, one
can obtain an impression of the density of the vegetation.
Using these ingenious pollen diagrams, Iversen and his collaborators
Earth-Sci. Rev., 2 (1966) 277-316

298 A.A. MANTEN
could show that the Allerod oscillation was characterized by a time in which
the forest temporarily closed without an immigration of thermophilous trees.
The oscillation (now preferably called an interstadial) was followed by a time with
a rather open park tundra, until in the Preboreal the forest definitely closed and
the Holocene Period began.
The concept of a park tundra or park landscape was introduced by Iversen
to denote a grass-sedge tundra with scattered stands of dwarf birches. Alongside
dwarf species of Salix and the arctic genera Dryas, Oxyria, Armeria, Empetrum
and Selaginella, there were also the genera Artemisia, Rumex, Thalictrum, Hip-
pophae and Helianthemum. The latter suggest a comparison with the central
European alpine, rather than with northern arctic, conditions.
From his pollen diagrams, IVERSEn (1947) also came to the conclusion
that once before in the Late Glacial, a temporary improvement in climate must
have taken place. He called this the Bolling oscillation. This interstadial repre-
sented a time with a park landscape, which was preceded and followed by a time
with a tundra vegetation. The following subdivision of the Late Glacial was thus
established: Oldest Dryas Time--Boiling Interstadial--Older Dryas Time--Allerod
lnterstadial--Younger Dryas Time. The latter is succeeded by the first stage of
the Holocene, the Preboreal, in which the first thermophilous trees appear.
This definitely meant the end of the earlier belief that the Late Glacial in
northwestern Europe was characterized by the simple succession of times with a
tundra vegetation, a park landscape and a subarctic forest. Later palynological
studies of Late Glacial material from the Mediterranean area, such as by MEN~N-
DEZ AMOR and FLORSCrtt)tZ (1962) on samples from Spain, revealed that a simul-
taneous plant-geographical zonation during the Pleniglacial did not exist either.
The most recent development in palynological studies of the Late Glacial
is the evidence presented by VAN OER HAmr~EN and VO6~L (1966), from Colombia,
Kenya, Spain and France, for an interstadial which is still older than the Bolling
Interstadial, and which they named the Susac/l lnterstadial. DREIMANIS (1966)
showed that this interstadial is of world-wide significance, being contemporaneous
with an interstadial described earlier from the Great Lakes region of North America,
with the Raunis Interstadial in the eastern Baltic area and the Plyusna Inter-
stadial in northwestern Russia.
Studies like those by Iversen caused a revival of interest in Quaternary
palynology after World War II. Mikkelsen, stimulated by Iversen, dropped Von
Post's old idea that Corylus pollen should never be included in the pollen sum.
Iversen, in the first diagrams made according to his new method, had not included
Corylus in his total pollen sum either. Mikkelsen demonstrated that diagrams in
which Corylus had also been included in the pollen sum gave a more understandable
impression of forest history.
Earth-Sci. Rev., 2 (1966) 277-316

HALF A CENTURYOF MODERNPALYNOLOGY 299
Vegetation and climate

In 1960, IVERSEN published a brilliant paper about the Early Postglacial forest
development in Denmark, in which the relationships between vegetation and
climate are thoroughly discussed. He pointed out that it is important to distinguish
between "pioneer species" and "climax species" of trees. The pioneer species have
a high reproduction rate, set fruits at an early age, produce many seeds year by
year and show an effective seed dispersal. Consequently these species readily
appear wherever favourable new conditions for tree growth are created. On the
other hand, they are weak in competition with others as they require much light
and have a relatively short life. Typical pioneer trees are Salix, Populus tremula,
Betula, and Sorbus. Trees of the second group, the climax species, generally
have a poor reproduction rate and seed dispersal; they cannot keep pace with
trees of the pioneer type in the initial stage; on the other hand, they are tenacious
and they will invariably suppress and supersede the pioneer species in the long
run. They have a long life, attain a great height, can endure much and also provide
ample shade. Typical climax genera are Quercus, TlTia, Fagus, and Ulmus. Wherever
vegetation is not in equilibrium with its surroundings, succession will set in and
continue until stability is reached. The final phase of stability is called the climax
vegetation. The word "climax" does not mean that the climax species are better
adapted to the climate in which they grow than any other species that thrive and
reproduce there. But owing to their competitive character they may form stable
communities, and not only successional stages as in the case of the pioneer species.
A forest of climax trees may be superseded by pioneer trees only after forest
destruction (hurricane, forest fire, man).
When interpreting pollen diagrams of the Postglacial, it should be kept
in mind that forest successions are extremely slow. After the retreat of the land ice,
it took thermophilous trees many centuries to migrate from south of the Pyrenees
and Alps to Denmark, and in the case of Fagus and Carpinus even millenia.
IVERSEN (1960), therefore, concluded that one cannot attach any climatic signifi-
cance to the geochronological boundaries in the Postglacial and Early Holocene.
The improvement of climate occurred faster than the immigration of the ther-
mophilous trees. In the Early Boreal, for instance, non-arboreal pollen shows
that the climate in western Europe was even more favourable than at present,
although the tree vegetation was comparable with the present situation in Lap-
land. Only in the Atlantic period did the climax, the mixed deciduous forest,
establish itself. Changes that took place thereafter mainly concerned the arrival
of the slowest migrants (Fagus, Carpinus) and human activities.
IVERSEN (1960) also obtained important information about the composition

of the climax forest. He realized that in pollen diagrams from peat bogs found
in forests, the Alnus-pollen percentage is overrepresented because Alnus grows
in and around bogs. Iversen, therefore, sought a small bog with walls steep enough
Earth-Sci. Rev., 2 (1966) 277-316

300 A.A. MANTEN
to prevent the development of a peripheral vegetation of alder carr. Furthermore,

the bog preferably had to be overshadowed by tree tops to prevent the pollen
contribution of Tilia, which is insect-pollinated, being overshadowed by the
pollen of wind-pollinating trees. He found the tiny bog Hertugdalen (diameter
25 m) in the Grib forest in the Danish island of Sealand and in a pollen diagram
from this bog Tilia pollen in Atlantic time reached 118 ~o if the total contribution
of the wind-pollinating trees is taken as 100~o. This shows that at that time
the bog was surrounded by a forest in which Tilia was the major component.
According to Iversen, this was the case on the high grounds; on low-lying mineral
soils Quercus dominated and with a still higher ground-water level the mixed
oak forest was superseded by an alder (Alnus glutinosa) cart, provided that the
water is not completely stagnant. Similar results were obtained independent
of IVERSEN (1960) by JAYSSEN (1960) from the southeastern Netherlands.
Ice Age
Palynology also proved to be of great value in extending existing knowledge
of the overall chronology of the Pleistocene.
Before the beginning of the Ice Age, large parts of the Northern Hemisphere,
including Europe, were covered with the famous Sino-American flora. In its
main composition, this flora is still present nowadays in Japan, China and North
America.
During the Pleistocene, the advancing land ice forced this flora to move
southwards. This presented no serious problems in North America and Asia,
where after retreat of each glaciation the flora again recolonized the original areas
if the climatic conditions permitted it. In Europe, however, the situation was
different. The east-west orientated mountain ridges and the Mediterranean
presented barriers against which the southwards-moving thermophilous flora
was forced to halt. This meant that after each glacial the Sino-American element
was less well-represented in the European flora, and consequently made Europe
a very suitable continent for the palynological identification of deposits from
the successive interglacials. A complicating factor, however, was that at no one
place is there a complete Pleistocene succession. In order to obtain a good picture,
one had to combine scraps of information from various localities, and, moreover,
the deposits found in these localities had, in many cases, been deformed by
the direct and indirect agents of the glaciations and had been eroded by melt
water, rivers or dust storms. It required a fair dose of acumen to collect the many
necessary pieces of information and to produce a general, cohesive picture out
of them. Someone who certainly has this gift is ZAGWIJN (1957, 1960). He has
contributed significantly to the present knowledge that in the Pleistocene there
have been six glacials, separated by five interglacials. WOLDSTEDT(1958) deserves
the credit for having made a representative selection from the many local names
for the glacials and interglacials which had gradually been introduced in the lit-
Earth-Sci. Rev., 2 (1966) 277-316

erature, thus establishing an acceptable chronology for the whole of Europe.

Studies of various kinds have shown that climatic fluctuations have also
occurred in tropical areas, where they were characterized by an alternation of
wet and dry periods. Palynological data (e.g., MAARLEVELDand VAN DEN HAM-
MEN, 1959) suggest that the pluvials correlate with the glacials of the higher
latitudes.
Palynology in archaeology
In Danish pollen diagrams, at about the transition from the Atlantic to the Sub-
boreal period, the pollen percentages of Quercus, Tilia, Fraxinus, and Ulmus
undergo a distinct, though only temporary, decline. A contemporaneous transitory
increase takes place in the Betula-pollen frequencies, a more lasting one in those
of Alnus, and the pollen of Corylus shows a pronounced maximum. IVERSEN
(1941) was the first to realize that these phenomena could be attributed to forest
clearance with the help of axe and fire and undertaken by Neolithic man, followed
by a characteristic regeneration of the cleared area through the invasion of pri-
marily Betula, Alnus, and Corylus. Iversen's pollen-analytical conclusion was
supported by the occurrence of Neolithic remains and charcoal layers with the
onset of the vegetational changes and the low but constant frequency of the
pollen grains of cultivated cereals.
IVE~SEN (1941) also pointed out that the pollen of Plantago major and
Plantago lanceolata act as landmarks in distinguishing the forest clearance by
European man from this Neolithic period onwards, and that the latter species prob-
ably came to Denmark with formal agriculture. GODWIN (1956) drew attention
to the fact that Plantago major, together with certain other heliophilous plants
which occurred in the open vegetation of the Late Glacial, reappear with the onset
of forest clearance as ruderals.
TROlLS-SMITH (1953, 1955, 1960) followed up an idea first launched by
FAEGRI (1944) that the decline of Hedera and Ulmus, which can also be observed
at the Atlantic-Subboreal transition might be attributed to their use as fodder
trees for stall-feeding of cattle, instead of to climatic changes, as IVERSEN (1941)
supposed. TroEls-Smith pointed out that in the Danish diagrams the pollen
of cultivated cereals, together with that of the ruderal Plantago, occurs in minute
traces before the forest clearance, but never before the Ulmus decline. The same
holds for Allium ursinum, which is also considered to be antropochorous. By suc-
ceeding in correlating the appearance of traces of cereal pollen in a pollen diagram
with the horizon of the A ceramics of BECKER (1948), which is considered to be
equivalent to the classical Ertebolle culture, TroEls-Smith and his associates
showed that the Ulmus decline did indeed take place at a time when prehistoric
man combined hunting and fishing with primitive agriculture. IVERSEN (1960)
Earth-Sci. Rev., 2 (1966) 277-316

302 A . A . MANTEN
seriously contested this explanation of the major decrease in the frequency of

Uhnus pollen and still feels that this is primarily climatically conditioned.
For a forest clearance by man, IVERSEN (1941) introduced the Danish term
"landnam". When this phenomenon was also recognized in other countries,
this word became internationally accepted in palynological and archaeological
literature.
A typical Dutch contribution to the application of palynology is the pollen-
analytical study of grave monuments, particularly by workers of the Institute
for Biological Archaeology of the State University at Groningen (A. E. van
Giffen, H. T. Waterbolk, W. van Zeist). Thus, spectra from Neolithic grave
monuments left by the people of the Funnel Beaker and Bell Beaker cultures
corroborate the type of cattle farming described by Trolls-Smith, whereas the
agricultural practices of the Protruding Foot Beaker culture correspond to the
so-called landnam of Iversen. A detailed review of this and similar subjects is given
by VAN ZEIST (1967).
Studies like those indicated above have greatly increased the importance
of palynology for archaeology. They have largely replaced the palynological
dating of archaeological objects, which lost most of its importance through
the discovery by W. F. Libby and his co-workers, in 1947, of the radioac-
tivity of 14C and its use as a method of dating. On the other hand, this radio-
active-dating method made it possible to establish accurately with which part
of the pollen diagram of a site an archaeological culture corresponds. This further
improved the use of pollen analysis in obtaining information about the envi-
ronment of prehistoric man.
Predominance of Cenozoic palynology
If we look at the present output in palynological literature, it appears that no

less than two-thirds of this is in the field of palaeopalynology. Some specialists
believe that the amount of palynological information unreleased in petroleum-
company files may exceed the amount now available in all the published literature
(CROSS, 1964, p.5). These two facts together lead to the conclusion that pre-
sumably over 80 ~o of all present-day palynological activities are concerned with
palaeopalynology. The geological-palaeobotanical impact on palynology is very
great indeed.
Within palaeopalynology the study of Cenozoic deposits still takes a
predominant place, although as was said before, the amount of pre-Cenozoic
palynological work is now rapidly increasing. Over 6 0 ~ of palaeopalynological
literature published in the first half of the nineteen-sixties or over 40~o of the
total palynological literature dealt with the Cenozoic, with a ratio Tertiary:
Quaternary of roughly 1:3.
Whereas initially the Cenozoic history of the earth was studied primarily
Earth-Sci. Rev., 2 (1966) 277-316

by geologists and palaeontologists, after World War II other, more modern

approaches to earth history--geochemical, geophysical--also contributed to
an increasing extent to an understanding of the events during that era. If a yearly
ratio modern/traditional is determined for the scientific publications dealing
with the Cenozoic, one finds a relative decrease in the contribution from the tra-
ditional disciplines. It is interesting to note, however, that despite this, phyto-
palaeontology appears to have maintained a rather constant 2 0 ~ of the total
literature on that part of earth history. This means that it has gained in importance
compared to zoopalaeontology and other classical disciplines. No doubt this
is largely due to the continuously increasing importance of palynology (MANTEN,
1965c).
Extraterrestrial life
In 1961, the Americans CLAUS and NAGY found remarkable particles in a carbon-
aceous meteorite. They called these "organized elements" and could find no
explanation for these, other than their being remains of once living micro-
organisms.
Palynology became involved in this controversial subject, as it was felt
that this is the branch of science which should be able to throw some light on
the nature of the organized elements. Since the subject has recently been reviewed
in this periodical (MANTEN, 1966), readers are referred for further information
to that paper.
Pollen morphology
As we have seen, initially the interests of pollen analysts working with Quaternary
material was predominantly directed towards the arboreal pollen. For plant-
geographical and plant-sociological reasons, however, attention was gradually
being given also to the pollen grains of other plants, as these occurred in the samples.
In turn, this made it possible to draw conclusions, from the collected pollen-
analytical data, of a palaeoecological, palaeoclimatological and other nature.
The idea was to be able to identify all pollen types which were encountered in a
particular deposit. For this purpose, much larger collections of pollen of recent
plants were required to enable comparisons of fossil material with pollen of
known botanical origin. All these pollen grains had to be accurately described,
with emphasis, of course, on the structure of the pollen wall. In this respect,
important work was done especially in the palynological laboratory of the Danish
Geological Survey, in Charlottenlund. A descriptive system was developed, which
initially was only intended for internal use. When, however, it was planned to
write a handbook of pollen analysis, Iversen decided to publish the terminology
([VERSEN and TROlLS-SMITH, 1950). In the book itself (FAEGRI and ]VERSEN,
Earth-Sci. Rev., 2 (1966) 277-316

304 A.A. MANTEN
1950), pollen morphology only has a moderate part. Nevertheless, the system
has had great influence on pollen morphology. The system was based on the
terminologies of Fritzsche, Wodehouse, and Erdtman.
After the appearance of ERDTMAN'S An Introduction to Pollen Analysis
(1943), this author has not done further work on the subject of pollen analysis.
Since then, he has concentrated entirely on pollen morphology. All plant groups
were systematically subjected to detailed studies. This led to the publication of
a two-volume book, of which the first volume (ERDTMAN, 1952) deals with the
pollen morphology of the angiospermous plants, and the second volume (ERoT-
MAN, 1957) with the pollen morphology of the gymnospermous plants, the ferns
(Pteridophyta) and the mosses (Bryophyta). Both volumes have become pioneering
works for further pollen-morphological researches all over the world. Many
plant morphologists, particularly those with a botanical background, found in-
spiration in these books for their own original studies.
It was said earlier that there has always been a clear relationship between
the development of pollen morphology and improvements in microscopical
techniques. After the introduction of the apochromatic lens, the next major
improvement came from the application of electron optics. Two kinds of electron
microscopical studies are to be distinguished: first, those using medium magnifi-
cation (magnification about 5,000), in which pioneering studies were carried
out by FERN~NDEZ-MoR/INand DAHL (1952) and MiJHLETHALER(1953); secondly
those utilising the utmost magnification available, in which important contri-
butions were made particularly by Erdtman and his school.
Palynological literature
Books
The books by Wodehouse, Bertsch and Erdtman, which have already been
mentioned, remained in use after World War II also. In addition to these, several
other important books have also appeared during the last two decades, including
the Textbook of Modern Pollen Analysis, by FAEGRI and IVERSEN (1950) and
Erdtman's two-volume work on pollen and spore morphology and plant taxonomy.
This work is especially valuable because of its great number of descriptions and
illustrations of pollen grains and spores from different families of plants, prepared
according to a very definite and exhaustive plan (ERDTMAN, 1952, 1957). In
between the two volumes, he also published a new edition of his introductory
book (ERDTMAY, 1954). In addition to various books devoted entirely to palynology,
there are several others in which a partial palynological content has been inte-
grated with other scientific information.
Journals
Mention has already been made of the Pollen Analys& Circular, of which 4 issues
Earth-Sci. Rev., 2 (1966) 277 316

were published in 1943 and an equal number in 1944. From 1945 onward, it
appeared under the name Pollen and Spore Circular, 3 issues being published in
1945, 2 in 1946, 1 in 1947, 2 in 1948, 1 in 1949 and the last one, no. 18, not until
1954. Then, the Circular became incorporated in the Micropaleontologist,
published by the American Museum of Natural History, New York. Soon there-
after, that journal was reorganized, to become, since 1955, the present Micro-
paleontology.
At present two international scientific journals are devoted solely to palynol-
ogy. These are Grana Palynologica, appearing since July 1954, under the eminent
editorship of Erdtman, and Pollen et Spores, issued since May 1959, under the
guidance of Madame Van Campo, by the Mus6um national d'Histoire naturelle
in Paris. A third periodical, Review of Palaeobotany and Palynology, which will
start publication early in 1967, includes in its scope not only the various fields
of palynological research, but also the study of plant macrofossils. In addition,
there are also some national journals and numerous other publications which
regularly or occasionally present palynological articles.
Bibliography
Palynology is rather unique in possessing an almost complete bibliography.
Since 1927, ERDTMANhas been publishing literature surveys in a series of papers
"Literature on pollen statistics", since 1932 with the addition "and related topics".
These appeared in Geologiska F6reningen i Stockholm F6rhandlingar. In Germany,
H. GAMS published bibliographies in the Zeitschrift fiir Gletseherkunde und
Glazialgeologie, also since 1927. When Word War II shut offthe Old World from
the New, the Pollen Analysis Circular took over this task for the North American
continent. The periodical went on publishing bibliographies throughout its exist-
ence. After the war, Erdtman resumed the work of compiling surveys of"literature
on palynology", in the years 1957-1959 for the subject of palaeopalynology
assisted by 3 issues of a Palynologie, bibliographie, issued by the Service d'lnfor-
mation g6ologique of the Bureau de Recherches g6ologiques, g6ophysiques et
mini6res, in Paris. Madame M. Van Campo deserves special mention by com-
piling a section "References bibliographiques" for Pollen et Spores, since 1959.
From 1960 onwards, she has been assisted in this by Madame N. Planchais,
from which year the section has been published as a bi-annual separate supplement
to that periodical.
International conferences on palynology
Palynological Conference during the Stockholm Botanical Congress

In 1950, the Seventh International Congress of Botany was held in Stockholm.
After the sixth congress had convened in Amsterdam, in 1935, the seventh congress
was originally scheduled to be held in Sweden in 1940. For obvious reasons, how-
Earth-Sci. Rev., 2 (1966) 277-316

306 A.A. MANTEN
ever, it had to be postponed. In 1946, the Swedish Organizing Committee decided

to invite a congress to be held in 1950. In connection with the preparations for
this congress, F. P. Jonker, of Utrecht, wrote a letter to G. Erdtman. In this letter,
which was dated July 9, 1949, Jonker wrote that both FlorschiJtz and he "regret
that, according to Communication No. 2 of the VII International Congress, there
is no section or subsection for palynology, and, as far as we can see from the
preliminary outline of the program, also no session on palynology in the section
of phytogeography. We especially regret this as Sweden is the classic country
of palynology. We wonder whether you are in a position to draw the attention
of the Executive Committee to this." Erdtman replied in his letter to Jonker of
August 2, 1949, that "Palynology will no doubt be adequately dealt with at the
forthcoming congress. I have brought your and Prof. Florschiitz' opinion as to
palynological conferences to the knowledge of the president of the congress, prof.
Carl Skottsberg." Preparations started for some sort of palynological meeting.
This palynological conference in Stockholm, 1950, was the first general
international gathering of palynologists. In his opening remarks of the first session,
on July 10, 1950, Erdtman mentioned that only a few weeks before the meeting,
the decision had been taken to organize a palynological conference in the same
way as the sectional meetings. It was then too late to incorporate all palynological
subjects in the palynological conference. Some subjects were, therefore, presented
in the section of experimental ecology and others in the section of phytogeog-
raphy. President of the Stockholm palynological conference was F. Firbas, and
vice-presidents were H. Godwin, H . A . Hyde and R. Potoni6. Four half-day
sessions were held on July 10 and 1 l, in which 6, 7 (plus a demonstration of a
soil sampler and two peat augers), 6 (plus a demonstration of the Palynological
Laboratory, by Erdtman) and 2 lectures were presented, respectively, and which
55, 50, 70 and 40 people attended. The first international palynological conference
was found to be very stimulating.
Palynological Conference during the Paris Botanical Congress

The Eighth International Congress of Botany, held in Paris in 1954, had a paly-
nological section in its programme from the moment its organization started.
The section was organized by Madame Van Campo and G. Erdtman was its
president; 34 lectures were presented.
It is interesting to note that the section, by means of a resolution, expressed
"the wish that palynologists working with industrial undertakings be permitted
to publish their material from a taxonomic point of view, without requiring that
all stratigraphical details be published."
Further meetings
The Paris congress remained the only one where palynology was represented with
its own special section at an international botanical congress.
Earth-Sci. Rev., 2 (1966) 277-316

In the nineteen-fifties national meetings also began to be held in various

countries, partly with restricted membership, a phenomenon which continued,
even more strongly, in the sixties.
First International Conference on Palynology

At the time of the Stockholm and Paris conferences, the rapid rise of pre-Quater-
nary palynology had just started. Lectures on studies of pre-Quaternary deposits
were presented at both conferences, but only constituted a subordinate part
of the total programme. In the next years, the older deposits, however, attracted
ever more interest and together with this the number of palynologists with a
geological background strongly increased. The development went together with
a rapid expansion of palynological activity in America.
Some of the American palynologists, notably those working at the Univer-
sity of Arizona, came to believe that international palynological conferences
should not solely be sections of botanical congresses. Among palynologists, there
are workers with geological, botanical, archaeological, medical and other back-
grounds. All of these should be enabled to meet each other and be stimulated
to cooperate. They obtained the support of many other North American palynolog-
ists and convened the official First International Conference on Palynology.
This was held in Tucson, Arizona, in April, 1962. The president was T. L. Smiley.
The conference was a significant event for this rapidly developing field of science
and the excellent programme was very successful. It is unfortunate that no volume
of proceedings, containing in full all the papers presented at this important
palynological event, has been published. Abstracts of the contributions to the
conference were published in Pollen et Spores, in 1962.
During the Tucson conference, an evening meeting was held, under the
chairmanship of N. F. Hughes, to discuss international cooperation. The desire
to have a second international conference was unanimously expressed. An Inter-
national Committee was established to arrange for the next conference. F. L.
Staplin was elected president, and as members N. A. Bolchovitina, M. Van
Campo, G. K. Guennel, F. N. Hughes, F. P. Jonker, R. M. Kosanke, G. O.W.
Kremp, L. A. Kuprianova. This committee later coopted as further members
G. von der Brelie, H.A. Hyde, S. Macko, E. M. van Zinderen Bakker and B. S.
Venkatachala.
Second International Conference on Palynology

It was initially hoped that the second conference could be held somewhere in
the U.S.S.R. However, an official invitation failed to come. Then, in 1963, Staplin
suggested Utrecht as location for the conference. Some time was allowed for
suggestions of alternatives, but as none was proposed, Utrecht was officially
nominated in Staplin's circular letter to the International Committee of January
Earth-Sci. Rev., 2 (1966) 277-316

308 A.A. MANTEN
7, 1964. The conference was held from August 29-September 3, 1966. A detailed
report of this meeting can be found elsewhere (MANTEN, 1967b).
Towards international organizations in palaeobotany and palynology
First attempt towards a palynological organization

In 1943 and 1944, U.S. pollen analysts discussed through their Pollen Analysis
Circular and other media, the desirability of establishing their own organization,
but at that time, they did not reach any positive decision.
The International Organization of Palaeobotany

On May 12, 1948, seven palaeobotanists met at the Palaeobotanical Department
of the Swedish Museum of Natural History in Stockholm. These were W. Gothan
(Berlin), B. Sahni (Lucknow), Isabel Cookson (Melbourne), and T. G. Halle,
G. Erdtman, O. Selling and Britta Lundblad (all from Stockholm); soon after-
wards they also received support from R. Florin (Stockholm). Together they dis-
cussed the question of whether it is possible to found an International Palaeo-
botanical Society for the chief purpose of issuing a Journal of Palaeobotany
(HALLE and SELLING, 1948).
At the 18th International Geological Congress, London, 1948, W . N .
Edwards and W. N. Croft organized on August 23, a palaeobotanists luncheon,
which was attended by some twenty scientists, among whom a number of palynolog-
ists. Among other topics, the foundation of an International Palaeobotanical
Society was informally discussed. The following resolution was then passed and
sent to Professor Halle: "That an informal gathering of palaeobotanists at the
18th International Geological Congress in London (1948) welcomes the idea
of an international palaeobotanical association, and hopes that concrete proposals
will be prepared for the Botanical Congress in Stockholm in 1950."
The eighth session of the Palaeobotany Section of the Seventh International
Congress of Botany, in Stockholm was devoted to the present position and needs
of palaeobotany. The meeting took place on July 19, 1950, under the chairman-
ship of H. Hamshaw Thomas, and was attended by 45 congress members. It was
decided that an International Assocation of Palaeobotanists should be established
to promote the advance of palaeobotany without interfering with existing insti-
tutions. To that end an organizing committee was formed, consisting of W. N.
Edwards (as convenor), H. N. Andrews, R. Florin, W. Gothan and R. Sitholey
as members.
It is worth mentioning here, with a view to the description of future devel-
opments, that at the Stockholm congress the two taxonomy sections (cryptogams
and phanerogams) adopted a resolution "that there be established an International
Association for Plant Taxonomy with a Bureau for Plant Nomenclature and
Taxonomy, and that the Association and Bureau include the International
Earth-Sci. Rev., 2 (1966) 277-36

Commission for Botanical Nomenclature, the Standing Committee for urgent

taxonomic needs and the International Commission for Plant Taxonomy".
The committee set up to prepare a palaeobotanical organization reported
to the Eighth International Congress of Botany, in Paris, in a meeting held on
July 12, 1954. At that meeting the official decision was taken to found an Organi-
sation Internationale des Pal6obotanistes (I.O.P.), which in its statutes is called
International Organization of Palaeobotany (cf. Congr. Intern. Botan., 8me,
Paris 1954, Compt. Rend. Sdances et Rapports et Communications, sections 3,
4, 5 et 6, pp.lll-121). The latter name became the official one. It was decided
that the I.O.P. should also represent the Subsection of Palaeobotany of the
International Union of Biological Sciences (which was newly established during
the Paris congress) and the Committee for Palaeobotany of the International
Association for Plant Taxonomy. The first president of the 1.O.P. was R. Florin
(Stockholm), vice-president Suzanne Leclercq (Li6ge) and secretary-general Ed.
Boureau (Paris).
The Paris discussions about a palynological organization

In 1950, F. Verdoorn, at that time secretary of the International Union of Bio-
logical Sciences, suggested the establishment of an international commission
on palynology under the I.U.B.S., with G. Erdtman, Stockholm, as chairman
ad interim. The activities of that commission remained mainly restricted to the
preparation of the journal Grana Palynologica and correspondence aimed at
cooperation and dispersal of information in the field of palynology.
In order to discuss the establishment of an international commission on
palynology on a broader basis, Erdtman and Madame Van Campo organized
an informal meeting in Paris, September 18, 1953.
A proposal was now made to the palynological section of the Eighth
International Congress of Botany, in Paris, 1954, to establish a permanent palyno-
logical section of the I.U.B.S., with Erdtman as president and Mine. Van Campo
as secretary. The proposal was discussed in a special session of the section with
Erdtman in the chair. This was a very uproarious meeting, as several palynologists
felt that for a proposal drawn up without prior consultation of wide palynological
circles, it went much too far. The proposal was voted down and the meeting
closed. To appease feelings, Roger Helm, President of the congress, convened
another meeting, under his chairmanship. It appeared then that several palynolog-
ists were not opposed to the idea of an international organization in itself. This
led to a new general session, with F. P. Jonker as chairman, to discuss the matter.
It was thought that the time was not yet ripe for an I.U.B.S. section, but some
kind of international cooperation was judged useful and desirable. A committee
was appointed, under P. B. Sears of Yale University, to investigate the matter
and to prepare a report. Nothing was ever heard from this committee.
Earth-ScL gev., 2 (1966) 277-316

310 A. A. MANTEN
Tucson and Utrecht

The matter of an international organization of palynologists was again raised
in a general meeting at the First International Conference on Palynology, held
in Tucson, Ariz., in 1962. With the events in Paris still fresh in every one's memory,
it was decided that the subject matter be handled carefully and that for the moment
activities should be restricted to the establishment of an International Committee
to ensure that a second international conference would be organized after a
number of years.
As it was to be expected that an international organization would become
a subject of discussion again at the second conference, upon his appointment,
the secretary of that conference undertook as one of his activities to explore
the problems and possibilities for such an organization. He worked from the
premise that if an organization would be judged necessary, then this would
have to be an overall association, that could act as the sole international re-
presentative of palynology and would embrace all fields of palynological activity.
The possibilities of affiliation with major international unions, as those for
biological sciences and geological sciences were also investigated, as well as co-
operation or a possible fusion with the International Organization of Palaeo-
botany. A working paper was presented to all participants in the Second Inter-
national Conference on Palynology, Utrecht (The Netherlands), 1966. The assem-
bly of that conference passed a resolution that this way towards the creation of
an international organization of palynologists should be further explored and
they elected a working group to carry out this task (see further MANTEN, 1967b).
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BROWN, C. A., 1957. Comments on European Tertiary pollen studies. Micropaleontology,

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3l6 A.A. MANTEN
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(Received November 7, 1966)
Earth-Sci. Rev., 2 (1966) 277-316

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Earth-Science Reviews - Elsevier Publishing C o m p a n y , A m s t e r d a m - Printed in The N e t h e r l a n d s

HALF A CENTURY OF MODERN PALYNOLOGY

Secretary and Editor, Second International Conference on Palynohz~y,

A short review of the history of palynology is presented. This branch of science

In 1966, palynology celebrates its 50th anniversary. Modern palynology is con-

Earth-Sci. Rev., 2 (1966) 277-316

POLLEN STUDIES BEFORE 1 9 1 6

Earth-Sci. Rev., 2 (1966) 277-316

Medical pollen studies

A specialized line of pollen studies developed in medicine, in relation to the

Earth-Sci. Rev., 2 (1966) 277-316

Observations of fossil pollen

Subdivision of the Postglacial and Holocene

For a correct understanding of the historical development of pollen analysis,

Earth-Sci., Rev., 2 (1966) 277-316

LENNART VON POST

Earth-Sci. Rev., 2 (1966) 277-316

THE RISE OF MODERN PALYNOLOGY

Quaternary pollen analysis

Earth-Si. Rev., 2 (1966) 277-316

Earth-Sci. Rev., 2 (1966) 277-316

Pollen analyses and archaeology

Palynology and coal geology

Earth-Sci. Rev., 2 (1966) 277-316

The nomenclature problem in Tertiary palynology

Earth-Sei. Rev., 2 (1966) 277-316

Palynology and the petroleum industry

Most petroleum companies initially showed great reluctance to make use of

Earth-Sci. Rev., 2 (1966) 277-316

Botanical microplankton and chitinozoans

Earth-Sei. Rev., 2 (1966) 277-316

thirties, when several European palaeontologists, among whom O. Wetzel, A. Ei-

Pollen studies in the service of medicine

It has already been mentioned that Scheppegrell revived the "gravity-slide"

Earth-Sci. Rev., 2 (1966) 277-316

Treatment of pollen allergies

Earth-Sei. Rev., 2 (1966) 277-316

AR~BRUSTER and JACOBS, 1934~1935), GRIEBEL (1930--1931), and especially by

After the great advances in pollen morphology made in the nineteenth

Earth-Sci. Rev., 2 (1966.) 277-316

First books on palynology

The first book on palynology appeared in 1935. It was WODEr~OUSE'Swell-known

A DISCUSSION ABOUT THE RIGHT NAME

Earth-Sci. Rev., 2 (1966) 277-316

statistical or morphological, or be they concerned with pollen-induced diseases

Earth-Sci. Rev., 2 (1966) 277-316

THE MATURITY OF PALYNOLOGY

Palynology and the petroleum industry

There is no doubt that it is the successful utilization of plant microfossils in

Earth-Sci. Rev., 2 (1966) 277-316

increasing applications of palynology in petroleum-exploration work in the

A critarehs, dino.flagellates and chitinozoans

Earth-Sci. Rev., 2 (1966) 277-316

trichospherous or otherwise) and other microplanktonic fossils (probably mostly

In 1946, IVERSEN published a new method of presenting pollen-statistical data.

Earth-Sci. Rev., 2 (1966) 277-316

Earth-Sci. Rev., 2 (1966) 277-316

Vegetation and climate