Wen 1962007001001

Wentia 7 (1-116)
Pollen morphology of the
euphorbiaceae with special reference
to taxonomy
W. Punt
Museum and
(Botanical Herbarium, Utrecht)
(received December 28th, 1961)
CONTENTS
Chapter I General Introduction 2
a. Introduction 2
b. Acknowledgements 2
Chapter II History
2
Pollen morphology
a. 2
b. Euphorbiaceae 4
Chapter III Material 5
Chapter IV Methods 6
a. Flowers 6
b. Pollen preparations 7
c. Preservation of pollen grains 7
d. Microscopes 8
e. Punched cards 8
f. Drawings 9
Chapter V Some nomenclatural remarks 9
Chapter VI Pollen morphology 10
Chapter VII Glossary 15
Chapter VIII Results 18
A. Pollen grains of the Euphorbiaceae 18
B. Discussion of the results

20
a. Phyllanthoideae 20
Antidesma configuration 20
Amanoa configuration 32
Phyllanthus nutans configuration 37
Breynia configuration 38
Aristogeitonia configuration 40
b. Crotonoideae 47
Croton configuration 47
Cnesmosa configuration 57
Dysopsis configuration 60
Plukenetia configuration 60
Chiropetalum configuration 65
Cephalomappa configuration 68
Sumbavia configuration 69
Bernardia configuration 73
Mallotus configuration 77
Claoxylon configuration 90
Cladogynos configuration 93
Hippomane configuration 95
Summary 106
References 107
Index
110
CHAPTER I
GENERAL INTRODUCTION
a. Introduction
have stated Lindau Wodehouse

Many investigators (e.g. 1895,
1935, Erdtman 1952), that pollen morphology can be of
great
for plant while it also known that in
importance taxonomy, was
of Erdtman
Euphorbiaceae several types pollen grains exist (e.g. 1952).
On the
suggestion of Professor Lanjouw, who himself has worked
on the Euphorbiaceae of Surinam, the author has investigated the pollen
of this of that From the result it
grains family area. was apparent that
in the Surinam Euphorbiaceae many
different pollen types could be
In the work the therefore extended

distinguished. present study was
to all the genera in the Euphorbiaceae, from which one or more species
have been examined. Besides of the
a description pollen grains
examined a
drawing of most of the pollen types is given. The
pollen
morphologic groupings found have been compared with the
systems
of several authors (Bentham and Hooker 1880, Pax and K. Hoffmann
1931, The taxonomic conclusions, however, have

etc.). can only a
character. Future investigators of the

Euphorbiaceae will have
provisional
to check the taxonomic notes and comments.
b. Acknowledgements
The have been carried the “Botanisch Museum

investigations out at
Herbarium”, Utrecht. The writer is indebted the

en
greatly to
director, Professor Dr. for him the

J. Lanjouw, giving opportunity to
the his institute and for his and

carry
out
investigations at
help
guidance in preparing this paper.
The author is very
much indebted
to Professor Dr. F. P.
Jonker who advised him in the special problems
of He the “Ministerie
pollen morphology. very gratefully acknowledges
Kunsten for the
van
Onderwijs, en
Wetenschappen” sponsoring
The author wishes his sincere thanks the
present study. to
express to
directors of the herbaria at Brussels, Leiden, Kew and Paris for the
and residence in
hospitality help during his these places. He is also
very grateful to the “Miquelfonds” which covered the expenses

of
his stay in London, Paris and Brussels where so much valuable material
could be examined.
CHAPTER II
HISTORY
a.
Pollen morphology
The study of pollen morphology started in 1675, when Malpighi
observed, that various types of pollen grains are to be found in different
He variations of colour and Grew

plants. especially saw shape. (1682)
of difference in He distinguished
was already aware a structure, too.
POLLEN MORPHOLOGY OF THE EUPHORBIACEAE 3
smooth and spiny pollen grains. In the centuries

next two
quite a
number of botanists enriched our knowledge of pollen morphology.;

In 1834 a
synopsis of the literature on the subject was
published by
Mohl, which contributed better of
v.
greatly to a
understanding
pollen morphology. He studied representatives of of the
most
plant
families and gave a descriptive classification of their forms, v. Mohl
that the of the the

recognized apertures pollen grains are most
features. Some later Fritzsche

important morphological years
published his principal work on pollen grains (Ueber den Pollen,

From the careful
1837). descriptions it is clear that, besides a study
of the
of the
shape and
apertures a pollen grain, structure was accu-
rately analysed. His drawing and description of Jatropha panduraefolia

gives an excellent idea of the pollen grain. The description even very
accurately reproduces the construction of the (see

croton-pattern
PI. I, E4 and PI. VIII, 1).
In the period after v. Mohl and Fritzsche we come across
publications with
drawings of
pollen grains as additions to
analytic;
of in Martius, Flora Brasiliensis). Most
drawings plants (e.g. botanists,
added but little C. A. H. Fischer
however, to
pollen morphology.
the first scientist in 1890,
was to give, a comprehensive comparative
of the data then available. From the of 2200 in
study study species
158 families he able draw, the conclusions:
was to
e.g. following
Pollen of related similar.

1: grains species are generally
2: Some families have more than a single basic form.
3: Sometimes unrelated have similar

plants pollen grains.
In 1883 Radlkofer made of obtain

use pollen morphology to a
better classification of the Acanthaceae. It \yas done more

profoundly by
Lindau in “Die natiirlichen Pfianzenfamilien”
(1895) by Engler and
Prantl when he stated: “Es ist deshalb nur wenn auf die
consequent,
Pollenbeschaffenheit die Einteilung der Acanthaceae gegriindet wird,
wie dies bereits von Radlkofer angcdeutet wurde und im folgenden
ganz streng durchgefiihrt werden soil”. It is obvious that, with regard

families, pollen had become wellknown and
to some
morpholqgy a
of the end of the 19th

indispensable part taxonomy at
century.
In
fact, a renewed general interest in pollen morphology emerged,
not from the side of botany, but from the side of geology. It was the
geologist v. Post who, in 1919, for the first time published a modern
diagram; the primitive pollen analysis had resulted

pollen analytic
in refined method in quaternary the
a geology. However, composition
of sound of the
a pollen diagram requires a knowledge morphology
of of the
pollen grains. Originally a
primary acquaintance pollen
grains of trees and anemophilous plants was considered to be
sufficient,
but and the of wider
more more
necessity a
knowledge of pollen types
became evident. Consequently the number of investigators who made
researches into the of taxonomic
pollen morphology special groups
increased.
Potoni£ in 1934 and Wodehouse in 1935 published works on
pollen morphology that formed the basis of our modern terminology.

4 w. PUNT
Potonie as well as Wodehouse needed a of well defined terms

system
for the details of
description of pollen grains. The construction and a
well
pollen grain were, however, not enough known to be arranged in
convenient
a
system. Nevertheless, many terms of these investigators
are still in use, although sometimes the circumscription of the terms
is more or less altered.
In 1950 Faegri and Iversen, and Iversen and Troels-Smith
published a
system on the morphology of pollen grains. Their ter-
excelled in exactitude and

minology simplicity. The well-defined terms
made it determine which in

quite possible to
pollen grains occur
their of
determination-key European pollen types.
At the same time Erdtman also worked at a of pollen
system
terminology. He published several papers on this subject, but his main
work and
(Pollen morphology plant taxonomy, Angiosperms) appeared
in 1952. In this book Erdtman extended of
proposed an
system
In however, the terminology
pollen terminology. many respects, seems
be of
too
complicated to
practical use.
Only pollen morphologists
with able all the mentioned
a large experience are to identify terms
in the
glossary. Moreover, the descriptions of terms are not
always as
exact as they should be. What, for instance, is the exact boundary
between brevicolpate and brevissimicolpate or foramen and fora-
minoid? Nevertheless, his book is of valuable data

a source
providing
information of the pollen of all the families in the
primary types
Angiosperms.
b. Euphorbiaceae
This It is
family is one of the largest in the Angiosperms. a difficult
one on account
of the many different species, so that it is not surprising,
few botanists of the whole
that only a possessed a general knowledge
family.
Since the present is work, the
report mainly a
pollen morphologic
lines will only briefly the authors
following comment on
principal
on Euphorbiaceae.
In the 19th A. de Jussieu, J. Mueller of Argau and
century
H. Baillon the In the
published important papers on
Euphorbiaceae.
of the Linnean Society (Botany) of 1880 Bentham reviewed
Journal
the work of Baillon and Mueller Arg. In the same paper
Bentham
extensive notes on his own system.

gave
1910-1924 Pax assisted Kathe
From (partly by Hoffmann)
monographed most of the genera in Euphorbiaceae in
Engler’s “Das
Pfianzenreich”. In the second edition of Engler and Die

Prantl,
Natiirlichen Pfianzenfamilien, Band 19c (1931), a revised synopsis
of 1931
of the Euphorbiaceae is given. In the present paper
this synopsis
has been used the basic classification of the
as family.
After Pax and K. Hoffmann only incidental remarks have been
made on the system of the Euphorbiaceae. Most of them were given in

floras of certain Leandri, Flora of
papers concerning areas (e.g.
Flora of Indo-Chine; Flora of
Madagascar; Gagnepain, Leonard,
Important improvements have been made by Croizat
Congo). on
several Schultes discussed the Hevea and its

genera. amply genus
allies. A very
important work is Webster’s publication on
Phyllanthus.
In his “The West Indian
monographic study species of Phyllanthus"
he used pollen morphology as one of the main characters to divide the
genus Phyllanthus into subgenera and sections.
CHAPTER III
MATERIAL
Herbarium material obtained from the herbaria

following was
used in the study;

London (BM): British Museum (Natural History).
Bruxelles (BR); Jardin Botanique de 1’Etat.
Bruxelles (BRLU): Herbier d’Afrique de l’Universit6 de Bruxelles.
Kew (K); The Herbarium, Royal Botanic Gardens.
Leiden (L): Rijksherbarium.

Paris (P): Museum National d’Histoire Naturelle Laboratoire de
Phanerogamic.
Utrecht (U): Botanical Museum and Herbarium.
Wageningen (WAG): Laboratory for Plant Taxonomy and Plant Geography.
the of has been avoided. in

Generally use
type-material Only
those cases, where no other specimens were available while flowers
has been used for

were present, type-material pollen investigation.
The classification of the Pax and K.
Euphorbiaceae as
given by
Hoffmann in Engler and Prantl, Die natiirlichen Pflanzenfamilien
ed. 2, Band 19c (1931), was the principal system used for the choice of
genera.
Besides this system attention has been paid to the new genera
after listed in
published 1931 and the Index Kewensis. As far as
possible all known genera of the Euphorbiaceae were examined. In a
number of genera it for various reasons,

was, impossible to
study the
fall in the
pollen grains. They following groups:
1. Male flowers unknown.
Croizatia Steyermark
Pax K. Hoffmann
Lasiochlamys et
Kuhlman
Paradrypetes
Phyllanoa Croizat
2. The material studied did contain male flowers,

not or
only a
few, or unripe ones.
Leandri
Claoxylopsis
Calicopeplus Planghon
Cometia Thouars ex Baillon
Corythea Watson
Ditta Grisebach
Keyodendron Leandri
Mettenia Grisebach
Monadenium Pax
6 w. PUNT
Pax K. Hoffmann
Neomphalea et
Neotrigonostemon Pax et K. Hoffmann
Ramelia Baillon
Hemsley
Riseleya
Schumacher Lauterbach
Syndyophyll um et
Sphyranthera Hooker, J. D.
Thelypetalum Gagnepain
3. Genera of which no material has been seen.
Acalyphopsis Pax et K. Hoffmann
Pax K. Hoffmann
Annesijoa et
Chlamydojatropha Pax et K. Hoffmann
Clarorivinia Pax et K. Hoffmann
Gitaria Pax et K. Hoffmann
Senefelderopsis Steyermark
On of the number of and the author

account large genera species
has found it be of the (both
impossible to
always sure
accuracy
taxonomic and nomenclatural) of the names
used in the
present paper.
Future studies will be

monographic perhaps prove some names to
incorrect. With all the descriptions of pollen grains, however, the
collector’s name and number and the herbarium, where the specimen
is In those when the has be
preserved, are given. cases name to
corrected the of the thus be

description pollen grain can always
placed under the right name.
In the Leiden a
Rijksherbarium at large collection of plants of
D’Alleizette is These plants, however, have not been collected
present.
by D’Alleizette himself. In the herbarium at Paris D’Alleizette
obtained from several collectors. These duplicates

duplicates were sent
to the Rijksherbarium without mention of the names and collectioning

numbers of the original collectors. As it lies beyond the scope
of this
work search for the of the the of

to names original collectors, name
D’Alleizette has been maintained in the work.

present
CHAPTER IV
METHODS
a.
Flowers
From each at least one species and when possible two species were examined.
genus
From those that can

be devided into two or more sections, species were
genera
chosen from different sections.
Usually the male inflorescenses of the Euphorbiaceae have many flowers, so that,
loss it flowers dissected.
generally speaking, no important occurs one or two are
-
Only those flowers of the Euphorbiaceae are useful that have just opened. When
pollen it is
the flowers are
younger
the has not yet ripened, so that impossible to
give an accurate description of shape and structure of the pollen grains. It is far
better to use old, outpollinated anthers than and undoubtedly unripe ones.
young
In the corners of their thecas the old anthers always have some pollen grains which,
by a special micro-method described below, can be isolated.
POLLEN MORPHOLOGY OF THE EUPHORBIACEAB 7
b. Pollen prepararions
For the study of the morphology of pollen grains it is desirable that cytoplasma
and intine are dissolved. The microscopic image becomes obscure and the important
cxine remains indistinct if intine and dissolved. For that
cytoplasma are not reason
Wodehouse’s methylene-green method, the lactic-acid method and

any
other
method, which does completely dissolve intine and considered

not cytoplasma are
less suitable.
The treatment of the pollen grains with the acetolysis-method of Erdtman (1943)
produces the best results. Everything inside the exine is dissolved, while, moreover,
a colouring of the exine takes place. This method, described by Erdtman in 1943
and 1952 for herbarium material, has, in its original form, a
great disadvantage.
Too much material is required to obtain a usable preparation. In order to reduce
this disadvantage a
micro-method (see below) has been applied, which leaves the
method as such untouched, but requires less material. If even only one
necessary
anther is sufficient.
Micro-method
The micro-method is a perfection of the method described by Willrath in a
publication of Potonie (1934).

A flower is boiled in water till it is free of air. Under a preparing-microscope
one anther (or more if the material allows) is dissected and laid in the pit of a
hollow slide. Some drops of acetolysis mixture (9 parts anhydric acetic acid 1
:
part
H2SO4 cone.) are dropped on the anther till the
pit has been filled with the mixture.
Now the slide is placed on a heating-apparatus as shown in Fig. 1. The end of the
heating-apparatus is heated by a
Bunsenburner. The flame should not be too high,
Fig. 1
but not too low either. The slide furthermore has to be removed so far from the
flame that the mixture cannot take fire. A of the mixture will
part evaporate.
When the slide has become almost dry some more drops are added. This is repeated
3 or
4 times till the remaining mixture is turning dark brown. When the preparation
once more is nearly “dry”, it is quickly cooled down by placing the slide on a stone
or iron surface. Under the dissecting microscope the pollen grains are to be seen
as brown globules. With anthers which have a solid skin, that cannot easily be
dissolved, it is desirable, while heating, to prick this skin in places with a needle
to make the acetolyse mixture

penetrate deeply enough into the anther.
c. Preservation of pollen grains
For the preservation of grains the method has been used.

paraffin-comprising
A small drop of glycerin (diam. 1 mm.) is laid on a slide. With a brush moisted in
glycerin the pollen grains are wiped off the slide or fished out of the liquid. In
8 w. PUNT
the drop of glycerin the pollen grains come off the brush more or less easily. This
is repeated till at least 10 grains are

present in the glycerin drop.
Now a little piece of glycerin-gelatin is added. The gelatin is melted and the
mixed with it. To the pollen grains from expanding abnormally

grains are
prevent
the whole should not be heated too much. It is of the utmost importance that the
grains can be studied from all sides, so that care should be taken that the cover-
glass does not flatten the grains. Therefore a granule of clay (no synthetic material)
is laid to the glycerin drop. The cover-glass is supported by the when
next now
clay
the preparation and the pollen grains cannot be compressed. By lightly
placed on
pressing the cover-glass the thickness of the preparation can be regulated. By

running melted paraffin under the cover-glass the grains are isolated from the air.
Webster (1956) rejects the method of sealing by paraffin on the ground of

up
the preparations not durable. In Utrecht, however, preparations ten old

years
are still in excellent condition. It is, however, essential that the layer of paraffin
should not be thin and the drop of glycerin should not the of
too approach edge
the cover-glass (Erdtman and Praolowski 1959).
Preparations made by the micro-method have the disadvantage that some acid
will enter with the pollen grains. This cause the colour to fade after some
may
time. After three however, form and structure were still unchanged. For
years,
that reason the macro-method of Erdtman is always preferable to the micro-method,

but the latter has the advantage that little material is sufficient, so that
very any
sacrifice of herbarium-material be prevented. In the of

rigorous may case flowers
with large anthers it is possible that of stamen suffices (Campanula,
even
part a
Lilium).
In describing pollen grains it is of
great importance that the method used for
preparation should be indicated, since the size of the pollen grain varies with it.
Though the expansion of a certain pollen grain is constant in either method, only
relative value can be attached to the indication of the size of that particular pollen
that have been treated in
grain. Only with regard to pollen grains precisely the
same the grain measures retain their value. Moreover, the size is influenced
way
by the grade of ripeness of the pollen. When a pollen grain is on the

verge
of
maturing,
form and structure
may already be fully developed, while the size has not yet
reached its maximum. It is, therefore, again emphasized that all absolute numbers
which are mentioned in the descriptions have a

relative value. For this reason
any
accurate calculation of sizes has been relinquished. The figures give an
average
impression of the size in these special circumstances and allow a mutual comparison
of the examined pollen grains.
The proportionate numbers, however, are more dependable. Proportions remain
the same in either method. Some proportionate numbers are, Polar Area
e.g..
Index (=P.A.I.) and Polar axis: Equatorial axis Pollen
(= P:E) (see Morphology
p. 12).
d. Microscopes
The pollen grains were studied with a Bausch and Lomb binocular (objective
97 x, oculair 10 X ), and an Olympus phase contrast microscope was used. The
employment of this microscope was found useful in detecting the configuration

very
of structure elements. Most of the also visible with the
structure patterns are ordinary
light-microscope, but, especially in the case of small structure elements, not without
difficulty. The phase contrast microscope immediately gives a

clear image of
the structure.
e. Punched cards
The of large of material methods of

study quantities requires special compiling
the knowledge obtained. A simple card-index is inadequate when there are
many
be the
genera
to mutually compared; great number of characters hampers
any
efficient A solution of this problem found in the use of punched
arrangement. was
cards. Punched card made for the well

systems were
pollenmorphologic as as
for the taxonomic characters. Thus particular botanical object could be

any
classified in two independent systems. The of within those
groups genera systems
are now mutually comparable so that it became possible to recognize correlations
of characters, which, without the punched cards, would have been found at the
cost of much time and labour or not at all.
A further advantage is that the investigator is compelled to look over the same
characters continually. Even the absence of a special character be of importance

may
and finds expression in this
system.
A disadvantage may
be that too much reliance is placed upon
the characters
noted. Often characters that are indicated in the same are not mutually
way
comparable. If stamens are this coalescence can take various forms.

e.g. connate,
There be investigators, who inclined to distrust this Nevertheless
may are
system.
the use of punched cards and other mechanic should undoubtedly be taken
systems
into consideration, especially when studying larger taxa. Surveying and work-
ing the continually increasing numbers of characters in a reasonable time

up
is becoming more and more difficult. Loss of time, chance of errors and the risk of
being incomplete to compel the taxonomist to utilize form of mechanic

are
apt some
Who would, at venture to revise the whole Solanum

taxonomy. present, genus or
Euphorbia? Only through a perfection and especially through an acceleration of
our methods the larger taxa will as

yet have a chance of being monographically
worked on.
f. Drawings
The pollen grains have been drawn in such a manner that in one drawing as
characters as possible are to be seen at the same time. Each drawing therefore
many
is midway between a scheme and a photographical reproduction. The pollen grains
have been drawn without a camera lucida or other drawing instruments.
Drawing has the advantage that the most important characters can be indicated
distinctly, that the easier distinguish from others. A disadvantage

so grains are to
is the subjective image which the drawing is to give.

apt
The other possibility of reproduction is making use of photography. For the
following reasons this method has not been applied;

1. In most cases
at least 4 photographs of perfect quality are required to obtain
a distinct picture of the grain.

2. In a photograph the structures of the pollen grains can never be pursued
distinctly, because the photographic image will only be sharp for a certain
optical section.
3. Deeper lying layers debase the photographic image and cause an insufficient
sharpness of the essential characters.
The plates have been drawn to scale. Each picture is 1000 X enlarged, except
some of the large pollen grains (e.g. Manihot and Croton matourensis),
,, which are
_____
less enlarged. In those cases

where the enlargement is not 1500 X the scale is
given in the text to the plates.
CHAPTER V
SOME NOMENCLATURAL REMARKS
Bischofia Blume
Although Blume dedicated this to Bischoff he spelled, in his “Bijdragen”

genus
the with single f. Obviously he latinised Bischoff to Bischofius.
(1826), name a
For this the of the should be written in the

reason name
genus original spelling.
Briedelia Willdenow
In his “Species Plantarum” (4:978. 1805) Willdenow deliberately spelled the
with i He commented follows the choice of the “Genus hoc

genus
_
e. as on name:
a Cluytia et Rhamno abunde diversum in honorem Clariss. S. E. Briedel nominavi.”
Although the name of the botanist in question is spelled Bridel it is not clearly
demonstrated that the author made an unintentional error. Under the rules of
the international code the should be written in the original spelling.
genus
10 w. PUNT
Cnesmosa Blume
In his “Bijdragen” (1825) Blume spelled this as Cnesmosa. In the Flora van
genus
Java, Praefatio in adnotation, he changed the probably for

“
name, grammatical
in Cenesmon. This is not allowed under the rules of nomenclature. The
reasons,
original spelling should be maintained.
Hasskarl
Koilodepas
Koilodepas was first published in 1855 in the Greek version (Versl. Med. Akad.
Wetenschappen Amsterdam 4: 139). Two later the altered in

years name was
Coelodepas (Flora 40; 531. 1857). As Croizat and Airy-Shaw have

(1942b) (1960)
stated, the validity of the spelling Koilodepas is unquestionable.
Omalanthus A. de
Jussieu
The first spelling of this appeared in A. de Jussieu’s “De Euphorbiacearum”
genus
in 1824. In his “Conspectus” (1828) H. G. L. Reichenbach altered the name
in
Homalanthus. Of course the first spelling has to be maintained. Carumbium is a

syn-
of Omalanthus. Reinwardt published this name as a nomen nudum in Oken’s
onym
Isis 1823. So Omalanthus is the name first published and therefore valid.
Romanoa Trevisan
In 1824 A. de Jussieu published the Anabaena. De Bory, however, had

genus
given the name Anabaina to a new algae (Cyanophyta) in 1821, so that the
genus
name of A. de Jussieu is a homonym. In 1918 Pax et K. Hoffmann the
gave
substitute Anabaenella this did that
name to taxon. They not know, however, an
older name was available. In 1848 Trevisan had substituted Anabaena A. de Jussieu
by Romanoa (Alghe Coccotale If the is held from

p. 99). genus separate Plukenetia,
Romanoa is the first correct name for Anabaena A. de Jussieu.
CHAPTER VI
POLLEN MORPHOLOGY
In of
describing pollen grains the number terms
employed should
be small This the of Faegri and

as as possible. was starting point
Iversen in 1950. One of the advanced the
arguments they was
difficulty of giving an analytic description of a

pollen grain in exact
terms. The however, that resulted from their at

system, attempt
PLATE I.
A. 1. pori; 2. colpi; 3. prolate pollen grain; 4. oblate pollen grain; P =

Polar axis;
E =
Equatorial axis.
B. polar view; 1. triangular; 2. three lobed; 3. circular; 4. convex triangular;

S smallest distance between colpi ends; G S: G Polar
= two =
greatest breadth; =
Area Index.
C. 1. and colpus nudate; 2. and c. with a membrana granulata; 3.

porus p. p.
and c. with an operculum; 4. and c. with a (thickening a or thinning b

p. margo
of the ekt.); 4c. costae pori and costae colpi; 5a. with a vestibulum; 5b and c.
porus
composite aperture with an atrium.
D. 1. 2. circular 3.
colpus transversalis; colp. transv.; equatorial colp. transv.;
m = meridional axis of the e. equatorial axis of the colp.
cop. transv.; transv.;
o
=
os; 4. costa transversalis; 5. costa circularis; 6. costa equatorialis.
E. 1. tectum; 2. tectum perforatum; 3. reticulum; 4.
croton-pattern.
F. 1. 2. 3. 4. 5. 6.
psilate; scabrate; verrucate; gemmate; clavate; baculate;
7. echinate; 8. pila.
POLLEN OF THE EUPHORBIACEAE 11
MORPHOLOGY
PLATE I.
12 W. PUNT
and
simplicity was inadequate, amplification was necessary. Especially
when the amount of material to be described is considerable many
difficulties liable arise. The number of characters of

are to a pollen
grain is comparatively small and
they often appear in many transitional
forms. Yet and definition of characters is desirable.
a precise exact
Erdtman, who, in his study of the Angiosperms (1952), met with

these
difficulties, therefore enlarged the terminology considerably. He
tried to nominate every transitional form, each nuance, which could
be achieved only at the cost of exactitude and convenient arrangement.
Nevertheless will feel attracted

many pollen morphologists highly
towards his method.
For the of additional character in

use
pollen morphology as an
this is The obvious characters

taxonomy system too complicated. more
will suffice. For that be the

reason a
compromise seems to
right course
for the of diverse Without far

description pollen grains. going as as
Erdtman the author has attempted to use a terminology as

present
without loss of exactitude and
simple as was possible completeness.
Pollen
grains are
composed of three layers. The central part is the
The middle the
cytoplasma. layer is intine; the outer layer the exine.
The exine of pollen grains consists of a particularly resistant substance
of character: is resistant
a lipoid sporopollenin. Sporopollenin even more
the attack than cutin. The exine substance is

to
by micro-organisms
also highly resistant to all sorts of chemicals; it can not, or only slowly,
be attacked fluid KOH, concentrated H2SO4 fluoric
by or even
acid (HF).
The exine which can be devided in two
layers, the endexine and
the ektexine, supplies of the data for
most
necessary pollen description.
These data fall into three
groups:
1. Shape
2. Apertures
3. Structure
1.
Shape
If a
pollen grain is not spheroidal we can
distinguish two axis:
Polar axis (P) (PI. I, A3 and

a.
A4)
b. axis I, A3 and
Equatorial (E) (PI. A4)
In equatorial view show which will be
pollen grains a
shape
determined the P: E. this
easily now by calculating According to
relation we
distinguish the following classes (Erdtman 1952):
P E > 2
: : perprolate
1,33-2 :
prolate (PI. I, A3)
1,14-1,33: subprolate
1 —1,14: prolate spheroidal
1 : spheroidal
1 -0,88: oblate spheroidal
0,88-0,75: suboblate
0,75-0,50: oblate I, A4)

(PI.
< 0,50: peroblate
In the of
polar view circumference a pollen grain can be:
1. I,
triangular (PI. Bl)
2. convex triangular (PI. I, B4)
3. circular I,
(PI. B3)
three lobed I,
4. (PI. B2)
2. Apertures
In the of Iversen and Troels-Smith the
system principal classifi-
cation is made according to the occurence of pori (PL I,
apertures:
and
Al), colpi (PI. I, A2) colpi transversales (PI. I, Dl, 2, 3) (in composite
apertures only).
Real and real formed in the ektexine.
pori colpi are by thinnings
A colpus transversalis, on the other hand, by a
thinning in the endexine.
For that of the endexine and the ektexine have to
reason apertures
be described of each other.
independently
Pori can be distinguished from colpi by calculating the length:
breadth. If length: breadth < 2 the aperture is a porus although the
is circle. If breadth 2 the
shape not
necessarily a pure length: >
aperture is a
colpus. The endexine below the edges of colpi or
pori
be thickened. These thickenings are
called costae
colpi and costae
may
pori (PI. I, C4c).
The
colpus transversalis (PI. I, D) is an easily recognisable character.
It is a thinning of the endexine. Yet this
aperture has been inaccurately
Faegri and Iversen
described by many investigators. (1950) confuse
this with a real porus when the colpus transversalis has the
aperture
shape of a circle. Consequently, when using their key for the deter-
it is
mination of European pollen types, sometimes difficult to decide
if “transversal furrow” is
a “porus longitudinal elongated” or a
present. Erdtman calls colpi + colpi transversales composite apertures ”.

the transversalis he introduced the
For term colpus new term os
(Pi- ora). If this term is used in that sense the name will have to be
rejected as
being a synonym. It may, however, be maintained for that
part of the
colpus crosses transversalis
the colpus (PL I, D3). that
Colpi transversales stand perpendicular on the colpi. Usually they are

Sometimes fuse
boat-shaped. they together into one single aperture
along the
equator of the pollen grain (colpus transversalis equatorialis),
but it is also that the
possible colpus transversalis becomes isodiametric
(colpus transversalis circularis PL I, D2). The medial dimension of the
colpus transversalis is rarely larger than the equatorial one (Pleiostemon
type, p. 30). Edges of
colpi transversales can also have thickenings;
the of the transversalis called:
according to shape colpus they are
circulares, costae transversales and

costae costae equatoriales (PI. I, D4, 5, 6).
To express the proportions of the colpus transversalis in exact
figures,
the is added the of this character. In
quotient m: e to
description
this formula m is the meridional axis and e the equatorial axis of the
colpus transversalis (PI. I, Dl).

Iversen and Troels-Smith the
(1950) introduced term Polar Area
Index The P.A.I.

(P.A.I. ). is the ratio between the greatest distance
of the ends of the breadth of the
two colpi and greatest pollen grain
14 W. PUNT
the Equatorial axis. This character is of great value for the

usually
description of pollen grains.
3. Structure
The ektexine furnishes characters of great diagnostic importance.

The description of these characters meets, however, with considerable
difficulties, so that it is necessary to use a
system as simple as possible
and highly desirable to avoid confusing terms (e.g. sculpture, see below).
The ektexine is of elements. Different forms and
composed structure
arrangements of these elements cause several structure

types to occur.
The elements the endexine

most common structure occuring on are
pila (PI. I, F8). Each pilum consists of a swollen

apex' (caput) and a
cylindrical basal Pila

part ( collum; homologous columella). are strictly
found on the endexine. They may occur ordinate or inordinate. Ordinate
form reticulum PI. I,

arranged pila can regular figures (e.g. E3) or
stand in rows
striae). Frequently capita of
pila are connate or
(e.g.u x
fused. If the fusion is only laterally, and less of the surface is covered
the Pollen
by capita, we
speak of tegillate pollen grains. grains are
when the fused 80 of the surface

tectate capita cover % or more
(PI. I, El). In that case the

pila form a second membrane outside the
endexine. If the tectum is not

completely closed (fusion of the capita
than 80 less than 100 the membrane is
more
% and %) perforate a
tectum
perforatum (PI. I, E2).
On the of the elements of different
top tectum we
distinguish structure
shape. According to Faegri and Iversen they are called: baculae,
clavae, echinae, gemmae,

scabrae and verrucae (PI. I, F2 t/m 7) (for descrip-
tions see glossary). Psilate pollen grains have no structure elements on
the tectum
(PI. I, FI).
Structure elements the of the may also be
on
top tectum
arranged
in different If is
structure
types (reticulum, striae). a tectum
present,
of the columellae called intra-reticulum,
corresponding structure
types are
intra-striae. Sometimes columellae can give an impression of a reti-

“
culum. This is, however, in which each columella
a
pseudo-reticulum”
represents a “lumen” and the space between the columellae, the “muri”.
A real reticulum is formed several elements.

always by structure
In the study no difference is made between structure and

present
Potonie all elements outside the
sculpture. According to
(1934), tectum
form the sculpture of a pollen grain. Potonie tried to give sharp de-
finitions of these terms, but admitted at the same time, that structure
and sculpture pass

into each other. Faegri and Iversen (1950)
also stated, that and confuse. For that
structure sculpture are easy to
reason it seems better to speak of structure elements and structure only.

CHAPTER VII
GLOSSARY
Annulus (Iversen and Troels-Smith G 4a

1950) (PI. I; en b)
Area which surrounds and is distinguished by
a
porus a thickening or a thinning
of the ektexine; thus forming a prominent or depressed margin of the latter.
Aperture (Faegri and Iversen 1950)
Any weak performed part of the ektexine endexine of

or a pollen grain (e.g-
colpus, colpus transversalis).
Atrium (PL b
I; 5a, en c)
A cavity inside composite aperture caused between
a
by a separation two
layers of the exine.
Bacula pi. Baculae (Iversen and Troels-Smith (PI. I; F 6)

1950)
-
Structure elements with least dimension 1 in the

at one or larger shape
of rods. Occur on the tectum only.
Baculate (Iversen and Troels-Smith 1950)

Pollen with
grains provided baculae.
Caput -
pi. Capita (Erdtman 1952)
Upper part of a pilum.
Clava -
pi. Clavae (Iversen and Troels-Smith 1950) (PI. I; F 5)
Structure elements with least dimension 1 in the
at one
fi or larger shape
of clubs. Occur on the tectum only.
Clavate (Iversen and Troels-Smith 1950)

Pollen grains provided with clavae.
Collum (Erdtman 1952)

Lower of pilum; homologous with columella.
part a
Columella —
pi. Columellae (Iversen and Troels-Smith 1950)
Lower of pilum. Columellae bear the tectum. In 1956 Faegri used
part a
the term as a
for pilum.
synonym
Colporate (Erdtman 1945)

Pollen grains with composite apertures.
Colpus pi. Colpi (Erdtman 1943) (PI. I; A2 and

Cl)
-
Longitudinal apertures of the ektexine. breadth > 2.

Length :
Colpus transversalis (Erdtman 1943) (PI. I; D 1, 2, 3)
Aperture of the endexine perpendicular to the colpus. Appears in composite
apertures only.
Composite aperture (Erdtman (PI. I; D 1, 2, 3)

1952)
Ektexine combined with endexine
aperture an
aperture (apertures not con-
gruent).
Costa pi. Costae (Faegri and Iversen 1950) (PI. I; C 4 and 4, 5,

D 6)
-
Thickenings of the endexine.
Costae circulares (D5): Thickened edges of circular colpi trans\ ersales.

16 w. PUNT
Costae colpi (C4c): Thickened endexine below the edge of colpi.

Costae equatoriales (D6): Thickened edges of colpi transversales extended along the
equator.
Costae pori Thickened endexine below the edge of pori.

(C4c):
Costae transversales (D5): Thickened edges of colpi transversales.
Croton-pattern (Erdtman 1952) (PI. I; E4)

Structure elements in rings of 5 or 6 around a circular area.
Echina -
pi. Echinae (Wodehouse 1928) (PI. I; F7)
Structure elements with at least one dimension 1 or larger in the shape

of spines. Occur on the tectum only.
Echinate (Wodehouse 1928)

Pollen grains provided with echinae.
Ellipsoid
with and short axis.
Inaperturate pollen grains a long a
Endexine (Erdtman 1943)

Inner of the exine.
part
Ektexine (Erdtman 1943)

Outer of the exine.
part
Equatorial axis (Van Zinderen Barker 1953) (PI. I; 3E and 4E)

The line perpendicular to the Polar axis in the equatorial plane.
Exine (Fritzsche 1837)

The usually resistant, layer of the wall of pollen grain.
outer, a
Gemma —
pi. Gemmae (Iversen and Troels-Smith 1950) (PI. I; F4)
Structure elements with at least one dimension 1 or larger of which the
lower constricted. Occur the tectum only.

parts are on
Gemmate (Iversen and Troels-Smith 1950)

Pollen grains provided with
gemmae.
Inaperturate (Iversen and Troels-Smith 1950)

Pollen grains without
apertures.
Intectate (Iversen and Troels-Smith 1950)
Capita of the pila covering less than 80 % of the surface.
Intra-reticulate (Faegri and Iversen 1950)

Columellae inside the tectum form a network.
Intra-striate (Faegri and Iversen 1950)

Columellae inside the tectum stand in rows.
Lumen -
pi. Lumina (Potonie 1934)
Spaces between the muri of a reticulum.
Margo (Iversen and Troels-Smith

1950)
Area which surrounds a colpus and is distinguished by a thickening or thinning
of the ektexine; thus forming a prominent or depressed margin of the latter.
Membrana granulata (Erdtman 1952) (PI. I; C2)
Colpus or
porus membrane (= endexine) with some scattered structure
elements.
Murus —
pi. Muri (Potoni6 1934)

Ridges separating the lumina of a reticulum.
Operculum (Wodehouse 1928; Faeori and Iversen 1950) (PI. I; C3)

Isolated of the ektexine which is separated from the edges of colpus
part a
or by a narrow zone in which the ektexine is missing or greatly reduced.

porus
Os —
pi. Ora (PI. I; D3, o)
That of a composite aperture which is formed by the crossing of the colpus
part
and the colpus transversalis.
Periporate (Iversen and Troels-Smith 1950)

Pori uniformly distributed the surface of pollen grain.
T over a
Pilum pi. Pila (Erdtman 1952) (PI. I;

—
F8)
Structure elements consisting of a swollen
apex (caput) and a cylindrical basal
(collum, columella). Occur the endexine only.

part on
Polar Area Index = P.A.I. (Iversen and Troels-Smith 1950) (PI. I; B3)
The of the distance between the ends of two furrows and
proportion greatest
the the the
greatest breadth of pollen grain (usually Equatorial axis).
Polar axis (Erdtman 1943) (PI. I; A3p and A4p)

The perpendicular line connecting the poles of a pollen grain.
pi. Pori (Erdtman 1943) (PI. I; A1 and

Porus -
Cl)
Circular of the ektexine. breadth 2.
apertures Length : <
Psilate (Wodehouse 1928) (PI. I; FI)

Surface of tectum smooth.
Reticulum (Potonie 1934) (PI. I; E3)

Structural
pattern in the form of a network in which columellae represent
the muri of the reticulum.
Scabra -
pi. scabrae (Iversen and Troels-Smith 1950) (PI. I; F2)

Structure elements visible but smaller than 1 Occur on the tectum only.
/i.
Scabrate (Iversen and Troels-Smith 1950)

Pollen grains provided with scabrae.
Sculpture (Potonie 1934)

A term proposed by Potonie for the joint structure elements outside the tectum
(p. 14).
Stephanocolpate (Iversen and Troels-Smith

1950)
Pollen grains with more than three colpi perpendicular to the equatorial plane.
(Iversen and Troels-Smith

Stephanocolporate 1950)
Pollen grains with more than three composite apertures.
(Iversen and Troels-Smith 1950)

Stephanoporate
Pollen grains with more than three in the equatorial plane.
apertures
Stria pi. Striae (Iversen and Troels-Smith

-
1950)
Structure elements standing in rows forming narrow grooves (Jt parallel).
Striate (Iversen and Troels-Smith 1950)

Pollen grains with striae.
18 w. punt
Structure
Form and of the structure elements.

arrangement
Structure elements
Individual elements either on the endexine or on the tectum.
Syncolpate (Iversen and Troels-Smith 1950)
Colpi anastomosing at the poles.
Tectate (Iversen and Troels-Smith 1950) (PI. I; El)

The of fused and together 80 of the surface
capita pila are cover
% or more
of a pollen grain.
Tectum (Iversen and Troels-Smith 1950) (PI. I; El)

The membrane outside the endexine formed by fused capita of pila.
Tectum perforatum (Iversen and Troels-Smith 1950) (PI. I; E2)

The tectum is not completely closed. Small holes are
present.
Tegillate (Erdtman 1952)

The capita of the pila are laterally united or fused and cover less than 80 %
of the surface of a pollen grain.
and Troels-Smith 1950)

Tricolpate (Iversen
Pollen grains with three colpi perpendicular to the equatorial plane.
Tricolporate (Iversen and Troels-Smith

1950)
Pollen grains with three composite apertures.
Triporate (Iversen and Troels-Smith 1950)

Pollen grains with three pori in the equatorial plane.
Verruca -
pi. Verrucae (Iversen and Troels-Smith 1950) (PI. I; F3)
Structure elements with at least one dimension 1 or larger in the shape of
/i
warts. Occur on the tectum only.
Verrucate (Iversen and Troels-Smith 1950)

with
Pollen grains provided verrucae.
Vestibulum (Iversen and Troels-Smith I; C5a,

1950) (PI. b)
Cavity inside a caused by a separation between two layers of the exine.
porus,
CHAPTER VIII
RESULTS
A. POLLEN GRAINS OF THE EUPHORBIACEAE
Pollen grains of the Euphorbiaceae show a number of types, which
can be distinguished more or less clearly. Generally pollen grains of

belong the pollen type. The dimen-
species inside one
genus
to same
but in the
sions and proportions may differ, principle pollen grains
are
identical. In some larger genera different pollen types occur in one
More often find that different genera

genus ( Tragia ,
Phyllanthus). we
the it is it is
possess same pollen type. Consequently not possible, as
with the in
Acanthaceae, to place a plant instantly a genus by means
of its pollen grains.

For the the pollen within the subfamilies
greater part types Phyllan-
thoideae and Crotonoideae be from each other.
can
distinguished clearly
Some however, show some affinities; Clutia (p. 77)
types, e.g. type
with Phyllanthus pentaphyllus subtype (p. 24), and Amanoa type (p. 33)
with Sumbavia type (p. 69).
According to the system of Iversen and Troels-Smith (1950) the
following principal types occur;
inaperturate, periporate, triporate, stephanoporate,

tricolpate, stephanocolpate, tricolporate, stephanocolporate.
Pollen grains with composite occur most frequently.

apertures
Especially the number of tricolporate pollen grains is large.
The varies from (P; E > 2,00) oblate
shape perprolate to
E In
(P : =
0,50-0,75). polar view the pollen grains are mostly convex
triangular. In some cases the circumference is triangular (d

Clutia type),
circular ( Moultonianthus type) or three lobed ( Omalanthus nutans
subtype).
The colpi are usually narrow,
but sometimes broad (Amanoa type,
Plukenetia In types the
type). many colpi are
accompanied by costae
colpi ( Antidesma type, Mallotus type).

The P.A.I. from
diverge large (> 0,7) to
syncolpate. Syncolpate
pollen grains are, however, rarely found in the Euphorbiaceae (Amperea
type)-
Colpi transversales, if present, can be of great diagnostical value.
Shape and size are characteristic for several pollen types ( Securinega
type, Hippomane type).
Pollen of often Sometimes the
grains Euphorbiaceae are tectate.
columellae the small that they

supporting tectum are so are
barely
visible (Alchornea type, Ricinus type). If the pollen grains are intectate,
they generally possess a reticulum. In a
few cases only the pollen grains
are intectate, pilate (some Phyllanthus species, Tragia fallax type). On
the tectum structure elements of different shape may be found;
e.g. clavae, baculae, echinae, etc.
The croton-pattern is a structure

type which we meet in many
genera
of the Crotonoideae. Outside the Euphorbiaceae this structure type
is found in Buxaceae and Thymelaeaceae (Erdtman 1952).
In the of the material the

following treatment investigated pollen
grains are first divided into the two subfamilies:
a. Phyllanthoideae
b. Crotonoideae
Inside these subfamilies the discussed

pollen grains are according
their in
to
similarity type.
Different in different So the
pollen grains are placed pollen types.
unit has been chosen as the base of the classification. Descriptions
type
of a are therefore as extensive as possible. If the differences
type are
20 w. PUNT
of minor the pollen grains in

importance, are
placed subtypes. Several
types can have some characters in common. To express the corre-
these assembled in After the

spondences, types are
configurations. diagno-
of the is the
ses pollen groups comment given on pollen morphologic
characters, mostly in their relation other Besides
to pollen groupings.
this comment in many cases a taxonomic discussion is added, in which
the value of in taxonomic is discussed.

pollen morphology problems
B. DISCUSSION OF THE RESULTS
a. Phyllanthoideae
Antidesma configuration
Tricolporate or stephanolcolporate (4). Pollen grains with a

Polar axis larger
than the Equatorial axis. P. rarely shorter than the E. (e.g. Phyllanthus acidus sub-
type).
Colpus transversalis usually with costae.
Pollen grains reticulate or not reticulate; never echinate.
The Antidesma is characterised

configuration by its prolate shape of
the
pollen grains. The Polar axis is
usually larger than the Equatorial
axis. In the the
Phyllanthus acidus subtype (p. 26) only pollen grains
are
spheroidal or
slightly oblate spheroidal. However, the pollen grains
resemble much the that it better
so Securinega subtype, seems to place
them in the same
type.
The three main of the the Antidesma type,
types configuration are
the and the Dicoelia type. The three types

Securinega type are
closely
related and have several characters in common e.g. shape, costae colpi
and costae transversales. The other types resemble the main
types
~~
but lack one of the important characters e.g. costae (Heywoodia type,
30) or have special characters (verrucae in the Zimmermannia
p.
type, p. 29).
Fig. 2
The differences between the subtypes are rather faint and difficult
to define and separate. The most reliable character for determination
and found in the

separation is colpus transversalis. These apertures
are bordered by costae which make them distinctly visible. In some
subtypes (e.g. Antidesma subtype) the rims of the costae transversales
run
parallel and the outer ends of the colpus transversalis are diffuse.
Most Phyllanthus species have costae transversales with rounded ends.
The ends of the transversalis in that distinct and

outer colpus are case
closed.
frequently even
Antidesma
type
Tricolporate; perprolate to prolate spheroidal.

Colpus transversalis small or narrow elongated; costae.
Colpi narrow;
costae colpi.
Tectate; psilate. Mostly not reticulate; rarely indistinct reticulate.
The Antidesma type differs from the Dicoelia type (p. 28) by the
In
shape and dimensions of the colpus transversalis. other characters
the two types cannot be differentiated.

From the the Antidesma type differs
Securinega type not only by the
of the transversalis but also the of the
shape colpus by structure
exine.
Taxonomic discussion
The be divided into

plants in the Antidesma type can two
groups.
The first combines the Antidesma with the Baccaurea
group subtype
and the second group includes the
subtype two Phyllanthus subtypes.
The first group comprises most of the genera which Pax and K.
united in the subtribe Antidesminae. The

Hoffmann (1922, 1931)
Antidesminae are chiefly characterised by the inflorescense which is
catkin-like in slender The characters of

or
spikes. following are
importance too;
1. The male flowers have few (six or less;

only usually five)
stamens round an ovarium rudiment (in Aporosa the ovarium rudiment
small
is or absent).
2. Usually a disc is present.
3. In the female flowers disc be the
a may present; styles are
usually bifid.
4. The plants are nearly almost dioecious. All the genera are
found in the Old World

except Hyeronima, which is the American
representative of Antidesma (Bentham, 1880).

Chonocentrum first Mueller as of
was recognised by a species Drypetes.
Pax and K. Hoffmann (1922, 1931) separated the plant from this
and it in the Pollen of the

genus placed vicinity of Discocarpus. grains
latter however different. Chonocentrum differs from
genus are
quite
the characters:
Discocarpus by following
1. Petals are never
present.
2. The stamens are not connate but free.
Some included in the Antidesminae Pax and K. Hoff-

genera by
mann possess pollen grains without an Antidesma type. As for the genera
Dicoelia, Richeria and Richeriella related Martretia

they belong to
types.
and have different
Hymenocardia totally pollen grains.
The second group of the Antidesma type includes many species of
Also of Reverchonia the

Phyllanthus. pollen grains possess Phyllanthus
In
pentaphyllus subtype. many respects the genus
is related to
Phyllanthus.
The of the however, is The male
branching plants, not
phyllanthoid.
disc, always present in' is rather indistinct. The
Phyllanthus, two stamens
are inserted on a thick disc-like tissue. As in Phyllanthus there is no
ovarium rudiment, while in the female flowers a disc is present.

22 w. PUNT
Antidesma subtype
Pollen grains tricolporate; perprolate -
prolate.
Colpus transversalis long; outer ends indistinct and diffuse. Edges of the costae
transversales parallel.
The pollen grains are strongly elongated and have narrow elongated
colpi transversales.
Though always referred to the genus

Securinega, Securinega congesta
has pollen grains largely different from the other pollen grains in the
genus.
The large P: E and narrow elongated colpus transversalis
the into the Antidesma

place pollen grains straight subtype.
Antidesma bunius (Linn.) Spreng. Antidesma subtype; perprolate.

Java [U] PI. II, 2 P =
37,5 E =
18,5 P:E =
2,04.
n n
m:e < 0,5 P.A.I. = 0,25.

Antidesma venosum Tul. Antidesma subtype; prolate.
Stolz 476 [U] P =
19 fi E = 13 P:E =
1,50.
n
Antidesma Elmer Antidesma subtype.
agusanense
Elmer 15881 Idem Antidesma

[U] venosum.
Antidesma diandrum (Roxb.) Roth Antidesma subtype.

Hohenacker 167 Idem Antidesma
[U] venosum.
Antidesma edule Merr. Antidesma subtype.

Ramos 1083 [U] Idem Antidesma venosum.
Antidesma ghesaembilla Gaertn. Antidesma subtype.

Hort. Calcutta 1897 [U] Idem Antidesma venosum.
Antidesma membranaceum Muell. Arc. Antidesma subtype.

Eels 126 [U] Idem Antidesma venosum.
Antidesma obovatum J. J. Smith Antidesma subtype.
Versteeg 1789 [U] Idem Antidesma venosum.
Hyeronima alchorneoides Allem. Antidesma subtype.

Krukoff 6110 P E P:E
[U] =
28,5 fi
=
12 n
=
2,20.
Hyeronima laxiflora (Tul.) Muell. Arg. Antidesma subtype.

Sandwith 544 [U] PI. II, 1
Thecacoris leptobotrya (Muell. Arg.) Antidesma subtype.

Brennon
Zenker 23 [U] P = 36 E =
17,5 P;E =
2,04.
n
Cyathogyne spathulifolia Pax Antidesma subtype.

Schlieben [P] P = 40 fi E =
20,5 P:E = 1,94.
fi
m:e± 0,5.
Cyathogyne viridis Muell Arg. Antidesma subtype.
F.H.I. 3936 [K] Too for measurements.
young
Apodiscus chevalieri Hutch. Antidesma subtype.

Pobequin 1909 [P] P =
30 n E =
19 fi P:E =
1,58.
Protomegabaria macrophylla Hutch. Antidesma subtype.
Andoh 5474 [K] PI. 3 P 27,5 E 19 P:E 1,45.
II, =
fi =
n =
Spondianthus preussii Engl. Antidesma subtype.

Versuchsanst. 164 [U] P =
26 E =
18,5 P;E =
1,40.
n n
Maesobotrya floribunda Benth. Antidesma subtype.

Lebrun 1039 P 21 E 15 P;E 1,40.
[U] =
iu =
/a
=
Maesobotrya dusenii (Pax) Hutch. Antidesma subtype.

Zenker 221 [U]
Securinega Muell. Arg. Antidesma subtype.
congesta
Black 2432 [U] P = 45 fi E = 26 n P:E =
1,73.
m:e < 0,5.
Baccaurea
subtype
Pollen grains tricolporate; subprolate -
prolate sphaeroidal.
Colpus transversalis small. Outer ends indistinct and diffuse.
Edges of the costae transversales parallel.

plate ii. 1. Hyeronima laxiflora; 2. Antidesma bunius; 3. Protomegabaria macrophylla;

5. Phyllanthus pentaphyllus; 6. Baccaurea
4. Aporosa lindleyana; sumatrana; 7. Reverchonia
niruri.
arenaria; 8. Margaritaria nobilis; 9. Phyllanthus
24 W. PUNT
In its exine this the Antidesma

structure
subtype is related to
subtype.
less and the
The
pollen grains, however, are elongated colpus trans-
versalis is smaller.
Baccaurea sumatrana (Miq,.) Muell. Arg. Baccaurea subtype; subprolate.

Hort. Bog. VI C. 185 A [U] PI. II, 6 P = 19 ft E = 15 ft P:E =
1,33.
P.A.I. = 0,35 —
0,3.
Baccaurea javanica (Blume) Muell. Arg. Baccaurea subtype.
Hort. 344 Idem Baccaurea sumatrana.
Bog. [U]
Baccaurea racemosa (Reinw. ex Blume) Baccaurea subtype.
Muell. Arg.
Hort. 553 Idem Baccaurea sumatrana.

Bog. [U]
Baccaurea Gagnep. Baccaurea subtype.
oxycarpa
(= Gatnaia annamica
Gagnep.)
P 21 E 16 P:E 1,31.
Eberhardt 3042 [P] =
ft
=
n
=
Aporosa lindleyana (Wight) Baill. Baccaurea subtype.

Hohenacker PI. 4 P 19 E 17,5 P;E 1,10.
352 [U] II, =
n
=
ft
=
P.A.I. =
0,2 —
0,25
dioica (Roxb.) Muell. Arg. Baccaurea subtype.
Aporosa
King 298 [U] Idem Aporosa lindleyana.
Aporosa frutescens Blume Baccaurea subtype.
Herbr Utrecht 022876 Idem Aporosa lindleyana.
Collector unknown.
Phyllanthus pentaphyllus subtype
Pollen grains tricolporate; prolate -

subprolate.
Colpus transversalis small; outer ends almost distinct.
Edges of the costae transversales rounded at the end.
The difference with the Antidesma subtype and Baccaurea subtype

is found in the
colpus transversalis.
There is also relation with the niruri subtype. The

a
Phyllanthus
latter however, is 4
subtype, —
colporate.
Phyllanthus pentaphyllus Wright ex Phyllanthus pentaphyllus subtype; prolate.

Grisebach P =
32 /I E =
22,5 n
P:E =
1,43.
Boldingh 7378 [U] PI. II, 5 m:e =
0,5 P.A.I. =
0,3.
Phyllanthus sublanatus Schum. et Thonn. Phyllanthus pentaphyllus subtype.
de Wit 1010 [WAG] P = 45 'n E =
30
n
P:E = 1,50.
P.A.I. =
0,45.
Phyllanthus stipulatis (rap.) Webster Phyllanthus pentaphyllus subtype.
Geyskes 13 [U] P =
24 ‘n E =
17,5 n
P:E = 1,37.
Phyllanthus amarus Schum. Phyllanthus pentaphyllus subtype.
B. W. 10 [U] P =
35,5V *E = 27 n P:E =
1,32.
de Wit 474 [WAG]
Reverchonia arenaria A. Gray Phyllanthus pentaphyllus subtype.
Warnock 10723 [U] PI. II, 7 P =
37 E'= 30 P:E = 1,60.
n
m:e =
ca. 0,4.
Savia erytroxyloides Griseb. Phyllanthus pentaphyllus subtype.
Wright 1434 [BM] P =
32 ii E*= 26 P:E = 1,24.
n
m:e < 0,5 P.A.I. =

0,25.
Phyllanthus niruri subtype

Pollen grains stephanocolporate (4); prolate.
Colpi transversales; costae transversales. Edges of the colpus transversalis
rounded at the end.
Except for the number of colpi the Phyllanthus niruri is in all

type
its characters identical with the Phyllanthus pentaphyllus subtype.
As in the columellaeare sometimes

Phyllanthus pentaphyllus subtype the
fairly coarse. The surface seems to form a reticulum. This is
pattern
however in niruri
a “pseudo-reticulum” (p. 14). Only Phyllanthus
real reticulum formed several columellae
a
by a
regular arrangement of
was seen. This reticulum was only to be seen with a phase-contrast
microscope and then very indistinctly.
Phyllanthus niruri Linn. Phyllanthus niruri subtype.

Fuertes 89 [U] PI. II, 9 P = 39 E =
21 P:E =
1,86.
n n
P.A.I. > 0,5. Indistinct intra-reticulate.
Phyllanthus guianensis Klotz. Phyllanthus nirwri subtype.

B.W. 4599 [U] P = 36
/z.
Phyllanthus hyssopifolioides H.B.K. Phyllanthus nirwri subtype.

Focke 1278 [U] P =
30 E = 18 P:E =
1,67.
n n
Phyllanthus urinaria Linn. Phyllanthus niruri subtype.
Lanjouw 246 [U] Too
young.
Chonocentrum cyathoforum (Muell. Arg.) Antidesma type-, tricolporate.

Pierre ex Pax et K. Hoffmann Colpus transversalis with costae.
Spruce 3781 [P] P:E > 1.
Too to give it a definite place

young
in the Antidesma
type.
Securinega type
Tricolporate or stephanocolporate (4); subprolate -

sphaeroidal (rarely oblate
sphaeroidal).
Colpus transversalis small, circular or broad elliptic; costae.
Colpi narrow;
costae colpi usually developed.
Intectate: distinctly reticulate; lumina 1-2 n-
Pollen grains small (smaller 30 usually 15—25 μ).

p,
This has small

type pollen grains with circular or broad
elliptic
colpi transversales. The reticulum is distinct and usually fairly coarse
in with the small

comparison pollen grains.
The Securinega comprises, besides the Securinega (and

type genus
of and the small
Flueggia), many species Phyllanthus genera Margarilaria
and Astrocasia. These genera are all
closely related (Webster, 1956).
The in Pax’s close
genus Aporosella, system placed to
Aporosa,
should be reduced section of Phyllanthus Webster
to a
according to
(1956). One of the principal reasons which led him to this conclusion
is the close relation acidus. In the
to
Phyllanthus fact, pollen grains
of Phyllanthus elsiae, belonging to the section Aporosella, are closely
related those of acidus
to Phyllanthus (p. 26).
Securinega subtype
Pollen grains subprolate -

prolate sphaeroidal.
Colpus transversalis circular or
broad elliptic.
Pollen the have

grains of Securinega subtype always a P: E > 1.
The of the transversalis is circular broad

shape colpus to elliptic.
Securinega suffruticosa (Pallas) Rehder Securinega subtype.

Cantonspark (Baarn) P =
22,5 E =
2\/j. P:E =
1,08.
/a
P.A.I. =
0,25 —
0,3. Lumina 1-2 fi.

26 w.punt
Securinega ramiflora (Aiton) Muell. Arg. Securinega subtype.

Gressitt P 23,5 £ 20/* P:E 1,17.
Linsley 229 [U] =
fi
= =
Securinega neopeltandra (Griseb.) Urban Securinega subtype.

ex Pax et K. Hoffmann
Shafer 12080 [U] P:E =
1,18 Lumina < 1 fi.
Securinega virosa (Roxb. ex Wiled.) Baill. Securinega subtype.

Reichs-Kolon. 506 [U] PI. Ill, 2
Margaritaria nobilis Linn. f. Securinega subtype.

Soeprata 2D [U] PI. II, 8 P =
22,5 n E =
19,5 P:E =
1,15.
/a
m:e > 0,5 P.A.I. = ca. 0,25.
Lumina ca. 1
/a.
Astrocasia tremula (Griseb.) Webster Securinega subtype.
Gaumer Sens 1261 P 29 E 25,5 P:E 1,14.
et [K] = =
n
=
m:e ca. 1 P.A.I. =

0,3.
Lumina 1-2 fi.
Phyllanthus reticulatus Poir. Securinega subtype.

de Wit 128 [WAG] P = 18 E =
16,5 P;E =
1,09.
n ft
Bakhuizen v. d. Brink 3540 [U] P.A.I. =

0,25. Lumina ca. 1
fi.
Phyllanthus discoideus Muell. Arg. Securinega subtype.

2789 P 22,5 E 18 P:E 1,25.
Leeuwenberg [U] =
n
=
n
=
Costae colpi. P.A.I. =

0,3.
Lumina 1-2
fi.
Phyllanthus capillaris Schum. et Thonn. Securinega subtype.

De Wit 229 [WAG] PI. Ill, 1 Stephanocolporate (4) or tricolporate.
P =
21,5 n E =
20,5 n P:E =
1,05.
P.A.I. = 0,5.
Phyllanthus acuminatissima C. B. Robb. Securinega subtype.
Elmer 15662 [U] P =
22,5 E = 18 n P:E = 1,25.
n
P.A.I. =
0,25. Lumina ca. 1

fi.
Richeriella gracilis (Merr.) Pax et Securinega subtype?
K. Hoffmann Too for reliable
young measurements,
Hyting 65438 [K] but colpus transversalis distinctly cir-
cular.
Phyllanthus acidus subtype
Pollen grains tricolporate; sphaeroidal -

oblate sphaeroidal.
Colpus transversalis circular.
Unlike the the

Securinega subtype pollen grains of the Phyllanthus
acidus sometimes oblate
subtype are
spheroidal or
slightly spheroidal.
The colpus transversalis is always circular. The exine structure is in
all the in the

respects same as
Securinega subtype.
Phyllanthus acidus (Linn.) Skeels Phyllanthis acidus subtype.

Versteeg 566 [U] P = E = 17,5 P:E = ca. 1.
n
P.A.I. =
0,2 —
0,3.
Phyllanthus muellerianus (O. Kuntze) Phyllanthus acidus subtype.
Exell Colpi short. Too
very young.
Zenker 253 [U]
Phyllanthus elsiae Urban Phyllanthus acidus subtype.
and Urban 4789 PI. 3 P E 20 P.A.I.
Krug [L] HI, = =
n.
> 0,5.
Richeria type
Tricolporate; subprolate.
Colpus transversalis narrow;
costae.
Colpi narrow.
Tectate (?); intra-reticulate. Lumina ca. 1 fi.
The Richeria is an intermediate between the Antidesma

type type
type and the Securinega type. The narrow elongated colpus transversalis
plate in. 1. Phyllanthus capillaris; 2. Securinega virosa; 3. Phyllanthus elsiae; 4. Drypetes

glauca; 5. Actephila excelsa; 6. Andrachne phyllanthoides.
28 w. PUNT
is of the Antidesma but the

typical type, distinct, rather coarse reticulum
is characteristic of the Securinega type.
Pax and K. Hoffmann well Bentham

(1931) as as (1880) placed
this in the Antidesminae. It occurs in the New World.
genus
Richeria laurifolia Baill. ex Muell. Arg. Richeria type; prolate.

Krukoff 8776 [U] P =
29 E =
24 P:E =
1,21.
fi
Lumina ca. 1
fx.
Dicoelia
type
Tricolporate; prolate —
subprolate.
Colpus transversalis broad and elongated to broadly elliptic; costae.
Colpi narrow;
costae colpi.
Tectate, psilate; sometimes intra-reticulate.
Pollen grains of medium size (ca. 25 -

50 /*).
fi
The Dicoelia elements of the Antidesma type as well

type possesses
of the Polar axis
as Securinega type. The pollen grains have always a
of least 25 /u; usually between 30 fi and 55 pt.

The
at
colpus trans-
versalis is large; broad and elongated or broadly elliptic, but never
circular.
Pax and K. Hoffmann (1922, 1931) included the genus Dicoelia

in the Antidesminae for its inflorescences. the
elongated However,
loose, often androgynous clusters along the rachis of the racemes is
different from inflorescence in the

any Phyllanthoideae (Bentham
1880).
Dicoelia subtype
Colpus transversalis broad and elongated; m:e < 0,4.

Usually not intra-reticulate.
Dicoelia affinis J. J. Smith Dicoelia subtype.

Hallier P 44 E 32 P:E 1,37.
1255 [U] = =
//
=
m:e < 0,4 P.A.I. =

0,3.
Indistinct reticulate.
Drypetes glauca Vahl Dicoelia subtype.

Sintenis 5111 PI. Ill, 4 P 34 E 27 P:E 1,26.
[U] =
n =
n =
m:e < 0,25 P.A.I. =

0,25.
Pax et Dicoelia
Drypetes macrophylla (Blume) subtype.
K. Hoffmann P = 45 E = 40 P:E =
1,16.
n n
Bogor VIII F. 48 [U] m:e < 0,4 P.A.I. =

0,35.
Intra-reticulate.
Drypetes laterifolia (Swartz) Krug et Dicoelia subtype.

Urban
Fuertes 824 [U]

Drypetes variabilis Uitt. Dicoelia subtype.
B.W. 6481 [U]
Andrachne subtype
Colpus transversalis broadly elliptic; m:e = ca. 0,5 or larger.

Usually intra-reticulate.
POLLEN MORPHOLOGY OF THE EUPHORBLACEAE 29
Andrachne phyllanthoides Mueix. Andrachne subtype.

(Nutt.)
Arc. P = 45 E =
35,5 P:E =
1,27.
/i n
Bush 14843 [U] PI. Ill, 6 m:e
=
0,5 P.A.I. =
0,2. Indistinct
costae colpi.
Andrachne telephioides Linn. Andrachne subtype.
Dalmatic 1937, 50 [U] Intra-reticulate.
Andrachne colchica Fisch. et Meyer Andrachne subtype.

Cult. Hort. Canton (Baarn) 4884 [U]
Andrachne aspera Spreng. Andrachne subtype.
Pappi 4892 [U] P = 43 ft E = 31 n P:E =
1,40.
m:e > 0,5. P.A.I. =
0,3.
Longitudinal striation.
Actephila excelsa (Dalz.) Muell. Arg. Andrachne subtype.

Hort. Bog. VII B. 271 [L] PI. Ill, 5 P 48 E 37 P:E
=
n
=
n
=
1,36.
P.A.I. =
0,3. Costae colpi.
Poranthera corymbosa Brong. Andrachne subtype.
Constable 30096 [U] P = 34 E =
29,5 P:E = 1,15.
n /j.
m:e ca. 0,5 P.A.I. =0,25.
Tectum perforatum; lumina of the
intra-reticulum 2-3
/i.
Poranthera microphylla Brong. Andrachne subtype.
Hastings [U] P = 24 ft
E =
19,5 n
P:E =
1,23.
Tectate; intra-reticulate; lumina ca.
1
p.
Zimmermannia type
Tricolporate; prolate spheroidal.

Colpus transversalis small; costae.
Colpi narrow.
Tectate; verrucate.
The the in the

Zimmermannia type is only type Phyllanthoideae with
verrucae. Except for this remarkable character the pollen grains
belong without doubt in the Antidesma configuration.
Zimmermannia capillepes Pax Zimmermannia type.

FI. Usambarica 5862 [BM] PI. IV, 1 P 58 E 51 P:E
=
/e
=
/e
=
1,12.
m:e > 0,5 P.A.I. = 0,25.
Leptonema type
Tricolporate; subprolate.
Colpus transversalis broad and large; costae narrow.
Colpi costae colpi absent.

narrow;
Tectate; psilate. The columellae are so small, that they are hardly visible.
The has relation with the Dicoelia

Leptonema type some type (p. 28).
The the It differs from the Antidesma
type misses, however, costae
colpi.
the broad and
type (p. 21) by elongated colpus transversalis.
Pax and K. Hoffmann in the subtribe Glochi-

(1922, 1931) placed
diinae Leptonema together with Glochidion and Breynia. It seems better
in the of
to keep Leptonema vicinity Phyllanthus and its allied genera,
stated Bentham
as
by (1880).
Leptonema venosa (Poir.) A. de Juss. Leptonema type.

D’Alleizette 6405 Mad. P =
27 E =
22,5 P:E =
1,21.
ju yu
Nov. 1905 [L] PI. IV, 2 m:e <0,5 P.A.I. =
0,5.
30 W. PUNT
Pleiostemon type

Colpus transversalis; m:e larger 1; 6-8
costae; m =
fi.
Colpi narrow; costae colpi. P.A.I. small. Pollen grains sometimes syncolpate.
Tectate; psilate.
The Pleiostemon the Antidesma

type belongs undoubtedly to con-
figuration. The
colpus transversalis however differs from all other
types by its meridional elongation. The m : e is larger than 1. More-
the Sometimes the

over, colpi are very long. pollen grains are even
the
syncolpate (colpi anastomosing at poles).
The this without doubt related

genera belonging to type are to
Phyllanthus and its allied genera. Mueller

(1866) reduced the genus
Pleiostemon section of Bentham

to a Phyllanthus and (1880) referred it
In the system of Pax and K. Hoffmann Pleioste-

to
Securinega. (1931)
is maintained close On
mon as a
genus
to Phyllanthus. the other hand,
is found in the subtribe in their
Lingelsheimia Drypetinae system.
Corresponding characters are:
1. In the male flowers an extrastaminal disc and an ovarium

present
rudiment wanting.
2. In the female flowers annular disc and bifid
an
styles present.
3. is Pleiostemon the male flower is sometimes
Calyx 6-partite (in
4-5-partite. Petals are absent.
4. Plants monoecious and found in Africa

only and
Madagascar
(.Dangyodrypetes ).
Pleiostemon verrucosus Sonder Pleiostemon type.
Cooper 312 PI. 3 P E P;E

[BM] IV, =
20 n = 19,5 fi = 1,03
m:e > 1 P.A.I. < 0,2.
Lingelsheimia frutescens Pax Pleiostemon
type.
Lebrun 3855 [K] PI. IV, 4 P =
22,5 E =
21 P:E =
1,07.
fi n
P.A.I. =
0,1.
Dangyodrypetes ambigua L£andri Pleiostemon
type.
Serv. Eaux et Forests 7675 [P] P=24 E =
22,5 P:E =
1,07.
n n
P.A.I. almost syncolpate-syncolpate.
Heywoodia type

Colpus transversalis broad; costae absent; m:e ca 1.
Colpi narrow; costae colpi absent.
Tectate; psilate. Columellae distinct.
As the Polar axis is larger than the Equatorial axis the type is placed
in the Antidesma There is, however, that
configuration. a possibility
Heywoodia is wrongly placed in this because of the
configuration
absence of costae.
Heywoodia lucens Sim Heywoodia type.

Bally 5133 [K] PI. IV, 5 P =
27,5 E = 25,5 P:E = 1,08.
n n
m:e ca. 1 P.A.I. =
0,3.
plate 1. Zi mmermannia capillepes; 2. Leptonema venosa; 3. Pleiostemon

iv. verrucosus;
4. Lingelsheimia fruticosa; 5. Heywoodia lucens; 6. Amanoa guianensis; 7. Pseudolachno-
stylis glauca.
32 w. PUNT
Amanoa configuration
Tricolporate-stephanocolporate (4-5). Pollen grains usually with a P. shorter
than the E.
Colpus transversalis usually with costae.
Pollen grains reticulate. Rarely not reticulate (some species of Amanoa and
the Discocarpus type).
includes
The Amanoa configuration pollen types with a Polar axis
shorter than the Equatorial axis and lesser than six (usually 3—4) colpi.
The grains reticulate in few These latter
pollen are
except a
species.
however, have characters in with related
species, so many common
reticulate species, that there is doubt that the

no
they belong to same
type-
.
As in the Antidesma the types difficult

configuration are to separate.
the Amanoa type and Savia related.
Especially type are
closely
The genera this all related.

belonging to configuration are closely
The division of the pollen grains into four types is not found again in
the of Bentham and Pax and K. Hoffmann.

systems Mueller,
Mueller removed Briedelia and Cleistanthus distinct
(1866) to a
subtribe of their valvate all the other

on account calyx (nearly genera
in have imbricate calyx). For this Pax and
Phyllanthoideae an reason
K. Hoffmann also these in tribe.

(1931) placed genera a special
Bentham took less of the imbricate and
(1880) account
calyx found,
in the much with Amanoa. So he
remaining characters, affinity placed
Briedelia and Cleistanthus in close The
proximity of Amanoa. pollen
grains of Cleistanthus cannot be separated from those of Savia. Except
for the striation the of the
longitudinal pollen grains Briedelia type
are
also close related to those of the Savia From these results
very type.
it is clear that morphology supports the of Bentham.
pollen statement
Bentham discussed in the of the Antidesminae.

(1880) Uapaca group
Pax and K. Hoffmann established subtribe for this
(1931) a
special
the the Savia it
genus. Although pollen grains belong to
type seems
better to
keep this genus
out of the subtribes with a Savia type. The
involucrum and the absence of a disc in the male flower are characters
the of any other genus

too
divergent for placing Uapaca in vicinity
with a Savia
type.
In the system of Pax and K. Hoffmann (1931) the
remaining genera
of the Amanoa divided into four subtribes. 1. Wielan-
configuration are
diinae. 2. Amanoinae. 3. 4.
Discocarpinae. Pseudolachnostylidinae.
In these subtribes the following genera
do not have the typical
characters of the Amanoa configuration.
1. Astrocasia. Webster the genus is related
According to
(1956) to
Securinega. The pollen grains undoubtedly belong to the Securinega
type.
2. The is classified with Amanoa by all
Actephila. genus together
authors. excelsa, however, of the
Actephila possesses pollen grains
Dicoelia Examination of the of other
type. pollen grains species
is necessary.
POLLEN MORPHOLOGY OF THE BUPHORBIACEAE 33
3. Chonocentrum. The of Chonocentrum without

pollen grains belongs
doubt to the Antidesma type (p. 21).
4. Savia Pollen
erytroxyloides. grains of this Savia species belong to the
Phyllanthus pentaphyllus subtype.
The have the

remaining genera following characters in common:
1. The calyx is 5-partite.

2. Frequently petals are present.
3. Male as well as female flowers have a disc.
4. In the male flowers an ovarium rudiment is

present.
5. Stamens are often connated into a column.
6. Seeds never have a caruncula.
Amanoa type
Tricolporate-stephanocolporate (4-5); suboblate-oblate spheroidal.

Colpus transversalis broad; ora large; costae usually present.
Colpi wide; no costae colpi.
Tectate or intectate; reticulate or not reticulate.
If the pollen grains are reticulate the lumina are at least 3 fi in diameter.
When the pollen grains are not reticulate the structure elements are
tall and
at least 4
fi.
Exine thick, at least 4
very /i.
The Amanoa is characterised reticulum and

type by a coarse a
thick exine.
The reticulum of the Amanoa type had to be considered as an intra-
reticulum since the muri are connected by a thin membrane, which

covers
the lumina.
Some species of Amanoa lack the coarse reticulum. Amanoa guianensis

has with echinae. The columellae
a tectum provided long baculae or
of Amanoa in and
grandiflora are, optical section, gemma shaped
inordinate In both Amanoa the exine is, however,
arranged. species
more than 4 thick.
fi
Amanoa oblongifolia Muell. Arg. Amanoa

type.
Krukoff 7015 [U] PI. V, 1 P =
33,5 n. E = 43 n P:E =
0,78.
Colpi wide. P.A.I. = 0,25. Lumina
up
to 11,5 fi.
Amanoa grandiflora Muell. Arg. Amanoa type.
Murca Pires 835 [U] P =
39 E =
45/i P:E =
0,87.
n
P.A.I. =
0,5. Not reticulate. Columel-
lae shaped, inordinate ar-

gemma
ranged.
Amanoa bracteosa Planch. Amanoa E 80
type. =
/i.
D’Alleizette 6392 Lumina to 10 P.A.I. 0,4.

[L] up fi.
=
Amanoa boiviniana Baill. Amanoa type.

D’Alleizette Mad. Nov. 1906 P 37,5 E 45 P:E 0,83.
[L] =
fi
=
n
=
P.A.I. =
0,4 —
0,5.
Lumina to 4
up fi.
Amanoa guianensis Aubl. Amanoa type.
Bafog S.F. 1192 [U] P = 46 E = 52 P:E =

0,88.
fi /i
Wood P.A.I. 0,3. Costae transversalis. Co-

Herb. 28 [U] PI. IV, 6 =
lumellaevery short. Baculate-echinate;

baculae to 7 Endexine thick.
up fi.
34 w. PUNT
Pentabrachion reticulatum (Muell. Arg.) Amanoa 3-4

type; colporate.
Jantzon P = 45 jit E =
48/i P;E =
0,94.
Zenker 2982 [L] Lumina to 5 P.A.I. =
0,4—0,5.
up ft.
Costae transversales.
Pseudolachnostylis glauca (Hiern) Hutch. Amanoa 2-3 colporate.

type;
Lanjouw 1181 [U] PI. IV, 7 P = 31 n E = 34
/i
P:E =
0,91.
Lumina to 4,5
up ft.
Savia
type
Tricolporate-stephanocolporate (4); subolate-prolate spheroidal.

Colpus transversalis usually narrow; ora
small. Rarely colpus transversalis
broad and ora large (Cleistanthus ferrugineus).

Colpi sometimes costae colpi present.
narrow;
Tectate or intectate; reticulate. Lumina of the reticulum 3 or smaller.
ft
Exine is medium thick, lesser then 4 but still distinct.
ft
In many respects the Savia type resembles the Amanoa type. The
lumina of the reticulum are, however, distinctly smaller. Since the
also smaller this difference is

pollen grains are relatively perhaps not
of fundamental besides the smaller the

importance, but, lumina, colpi
and the small, it better the
are narrow ora so seems to
separate pollen
the Savia from of the Amanoa
grains of type
..
those type.
_ .
Some with Savia

species a type (Cluytiandra madagascariensis, Lach-
have Polar axis than the axis.

nostylis hirta) a
larger Equatorial By
this character the pollen grains should be in the Antidesma
placed
The exine however, is different from
configuration. structure, quite
the Antidesma and is, the other full agreement
configuration on hand, in
with the Savia type.
Savia andringitrana Leandri Savia

type.
Decary 1926 [L] P = 26 E = 29 P:E =
0,88.
ft n
m;e < 0,5. P.A.I. =
0,4.
Colpi narrow and short. Lumina 3 ft.
Savia (Sw.) Wield. Savia
sessiliflora type.
Eggers 1905 [P] P =

24,5 E =
29 P:E =
0,85.
fi !x
P.A.I. =
0,3.
Blotia oblongifolia (Baill.) Leandri Savia
type.
D’Alleizette [L] P =
25,5 E =
28.5 P:E =
0,89.
n n
Mad. 4 Mai. 1918 P.A.I. =

0,45.
Colpi short and narrow. Lumina 1-2
ft.
Wielandia elegans Baill. Savia 3—4 colporate.
type.
D’Alleizette Seych. [L] P = 37 /i E = 41,5 P:E =
0,88.
n
P.A.I. =
0,65. Lumina = 1-2 /a.
Colpi narrow and short.
Muell. Arg. Savia prolate spheroidal.

Lachnostylis hirta (Linn, f.) type;
[= Discocarpus hirtus (Linn, f.) Pax et P =

37 E 35 P:E =
1,05.
n- —
fi
K. Hoffmann] P.A.I. =
0,3. Colpus transversalis with
Bolus 2396 [K] costae. Lumina 1-2 fi.
Cluytiandra madagascariensis Leandri Savia prolate spheroidal.

type;
D’Alleizette Mad. 6402 [L] P = 32 E = 31 P:E =
1,04.
fi n
P.A.I. =
0,3. Costae transversales. Re-
ticulum indistinct. Lumina 1-2

fi.
Cleistanthus winkleri Jabl. Savia E 32
type. =
ft.
IX C. 165 A. [U] Costae colpi.

Bogor
Cleistanthus dichotomus J. J. Smith Savia type.
Gjellerup 142 [U]
1. Amanoa oblongifolia; 2. Cleistanthus ferrugineus; 3.

plate v. Uapaca heudelotii;
4. Discocarpus essequeboensis; 5. Briedelia monoica; 6. Breynia fruticosa; 7. Breyniopsis
pierrei; 8. Glochidion sericeum.

36 w. PUNT
Cleistanthus ferrugineus Muell. Arg. Savia prolate.

type;
Hort. Bogor. IX C. 43 A. [U] PI. V, 2 P =
44 E =
33 P:E =
1,34.
P.A.I. = 0,4 —
0,5.
Colpus transversalis isodiametric; no
costae; ora large. Lumina 1-2 fi.
Uapaca heudelotii Baill. Savia oblate spheroidal.

type;
Zenker 59 [U] PI. V, 3 P =
22 E =
25 P:E =
0,88.
fi n
m:e 0,5. P.A.I. =
0,5. Tectate.
<
Uapaca kirkiana Muell. Arg. Savia type.

Stolz 622 [U] P =
27 E =
29,5 P:E =
0,93.
n
Tecta te.
Briedelia type
Tricolporate; oblate spheroidal.

Colpus transversalis broad; costae.
Colpi narrow at the ends, broadened in the equatorial part. Sometimes costae
colpi present.
Intectate; reticulate. Lumina small and in chains along the colpi (pseudo-
striate). Muri distinct but not tall.
This is related the Savia The difference

type closely to
type. greatest
is found in the longitudinal of the lumina of the reti-
arrangement
culum. The chains of lumina give the pollen grains a striate appearance,
which is, however, striation caused

only a
“pseudo-striation”: a not
but the lumina of the reticulum which

by individual columellae by
of course, of several columellae.
are, composed
Briedelia monoica (Lour.) Merr. Briedelia

type.
Endert E 1066 PI. V, 5 P 24 E 25,5 P:E 0,94.
[U] =
ju =
fi =
m:e > 0,5. P.A.I. =
0,35. Lumina
< 1
ft.
Briedelia glauca Blume Briedelia type. E = 31

/a.
Winckel 865 B [U]
Briedelia assamica D. Hook. Briedelia E =31 Lumina 1-2
J. type. /t. /i.
Herb. East Ind. Comp. 4890 [U]

Briedelia stipularis (Linn.) Blume Briedelia E 42
type. =
//.
v. Leeuwen-Reynvaan [U] m:e < 0,5. P.A.I. = 0,25.

Godefroya rotundata Gagnepain Briedelia type.
(Jabl.)
P 32 E 33,5 P:E 0,96.
Gourgaud [P] =
n = n =
P.A.I. = 0,25. Costae colpi.
Discocarpus type
Stephanocolporate (4-5); oblate spheroidal.

Colpus transversalis small; no costae present.
Colpi narrow. No costae colpi present.
Tectate, psilate. Not reticulate.
The Discocarpus type is an intermediate between the of

type types
the Amanoa and the
configuration types of the Antidesma configuration.
The small columellae and the absence of a reticulum are characters
in favor of placing the in the Antidesma configuration. On the

type
other hand the oblate shape, the absence of costae and the 4—5
colpi
are more in with the Amanoa configuration. For these latter
agreement
it better discuss the type in the Amanoa
reasons seems to
configuration.
TC
Discocarpus essequeboensis Discocarpus
T HT7SPH
type.
Schomburgk 659 [U] PI. V, 4 P =
34,5 ft E =
37/t P:E = 0,94.
Ora small; m:e > 0,5. P.A.I. =
0,6.
Phyllanthus nutans configuration
Pollen grains with an areolate structure.
After of Iversen, Erdtman recommended the

a proposal term
areolatus for the structure described under Phyllanthus nutans

type
The definition him is: “Pollen with small
(1947). given by grains
areas
separated by small grooves forming a
negative reticulum”.
Webster that the areolate
(1956) states pollen grains are mostly
found in the New World. Some of in the Old
species Phyllanthus
World, however, have areolate
pollen grains that are superficially very
similar to those of the New World.
Phyllanthus nutans type
Periporate; the surface of the pollen grains is divided into or hexa-

pentagons
The pori are situated in the angular points.
gons.
Pori small.
Intectate. From each two of pila lead to the other

aperture rows
apertures.
In the centre of the and hexagons the pila form irregular reticulum.
penta- an
Phyllantus nutans Sw. Phyllanthus nutans type.

Hooker, Jamaica [U] PI. VI, 2 Longer axis 27,5 Shorter axis 25
/r. /*.
ca. 25 pori.
Phyllanthus adianthoides Klotzsch Phyllanthus nutans type.

Hulk 21 [U] Diam. =
17
/s.
Phyllanthus speciosus Jacq,. Phyllanthus nutans type.

Fresh material, Greenhouse Utrecht Diam. = 20 Pori ca. 15.
/i.
Dendrophyllanthus type
Tricolporate oblate.
(?);
Colpus transversalis circular; costae.
Colpi narrow; operculum consisting of one row of pila.

very
Intectate; not reticulate.
Without doubt the pollen grains are related to the Phyllanthus nutans
type in their exine structure. The surface of the pollen grains is
divided into three of the

regions, separated by strings pila, “colpi”.
Each consists of three of pila. If the
“colpus” rows two outer rows
represent the borders of the colpus, the inner row must be taken as an
the space between the is that it

operculum. However, rows so small,
weak in the ektexine
can hardly be regarded as a performed part (see
definition aperture: Glossary). For this reason it is possible to defend
the that the of

opinion, pollen grains Dendrophyllanthus are triporate.
In that case the rows of pila would only be an ornamental peculiarity.
Nevertheless, the rows look so much like real colpi, that in the author’s
opinion it is better to maintain the pollen grains in the group of the
tricolporate.
Dendrophyllanthus ripicolus Guill. Dendrophyllanthus type.

Mackee 3629 [L] PI. VI, 3 P =
19,5 E=27 P:E =
0,72.
n n
m:e = ca. 1.
38 w. PUNT
Breynia configuration
Stephanocolporate (colpi sometimes with two ora).

Colpus transversalis circular rarely somewhat elliptical; costae.
Colpi narrow and long.

Tectate or
intecta te.
its exine the is related the

By structure Breynia configuration to
Antidesma Most however, have Polar

configuration. pollen grains, a
axis than the axis in the Amanoa

shorter Equatorial as configuration.
The of the number of
typical character colpi with often two circular
transversalis justify this configuration.

colpi per colpus seem to
Breynia type
Stephanocolporate (colpi diorate); oblate -

subprolate.
Usually two circular colpi transversales colpus, some colpi have only one
per
colpus transversalis; costae.
Colpi narrow;
sometimes costae colpi.
Intectate or reticulate or not reticulate; lumina small.
tectate;
The is the number of

Breynia type easily recognised by large colpi
and the double circular transversales. In the
(six or
more) colpi
species of Breynia one of the two colpi transversales is often reduced.
The remaining endexine aperture is never situated in the equatorial
towards the ends of the
plane but colpus.
Hitherto the genera belonging to this have never been placed

type
in author. Mueller the relation-
one group by any (1866) recognised
shared
ship of
Agyneia with Sauropus, which opinion was by Bentham
and Pax and K. Hoffmann

(1880) (1922, 1931). Breynia and Glochidion,
the other hand, are related
on
always distinguished as genera closely
to Phyllanthus. Glochidion is even often reduced to a section ofPhyllanthus.
and
Pax and K. Hoffmann
keep Glochidion, Breynia Breyniopsis in the
subtribe Glochidiinae. In the author’s opinion the above genera should
all be in (either subtribe other They

placed one group or
taxon).
have the following corresponding characters:
1. The number of is six sometimes

sepals usually (iGlochidion fewer).
2. In the male flowers generally three connated stamens form a
column. Connectives are frequently appendiculated. Anthers
extrors. There is no ovarium rudiment nor a disc. As Bentham
the scale-like the base of the of

states, thickenings at sepals
and be regarded disc glands.
Agyneia Sauropus are not to as
3. In the female flowers disc glands are wanting.

4. The monoecious and
plants are generally (Some Glochidion species
and in the Old World.
Arachnodes, however, dioecious) only occur
Breynia subtype
Pollen grains oblate spheroidal -
oblate.
Intectate; reticulate; lumina 1-2

//.
Breynia fruticosa (Linn.) Benth. Breynia subtype; oblate.
J. and M. S. Clemens P E P:E

3182 [U] =
21 fi
=
27 /x
=
0,76.
Beekler 2 [U] PI. V, 6 Colpi transversales situated almost at
the end of the colpi. Colpi not always

provided with two colpi transversales.
Colpi 8-10. Lumina 1-2 p.
Breynia racemosa (Blume) Muell. Arg. Breynia subtype; oblate.
Bakhuizen v.
d. Brink [U] 5-6 colpi.
Breynia nivosa (W. G. Smith) Small
Breynia subtype.
Collector unknown. 7-8 colpi.
Agyneia bacciformes (Linn.) A. de Jussieu Breynia subtype; oblate.
v. Leeuwen-Reynvaan 1978 [U] P =

27,5 E =
32,5 P:E =
0,85.
n /j.
Always two colpi transversales

present.
Golpi 5-6. Costae colpi.
Sauropus androgynis (Linn.) Merrill Breynia subtype; oblate spheroidal.
Doctors v. Leeuwen, Depok 7-5-’11 P =
25,5 iu E =
28 P:E =
0,90.
[U] 8-9 colpi.
Breyniopsis subtype
Pollen grains subprolate.

Tectate; not reticulate.
The pollen grains of Breyniopsis are related to the pollen grains of

the number of and the double circular
Breynia by colpi colpi trans-
versales. The shape and structure

are, however, quite different.
Breyniopsis pierrei Beille Breyniopsis subtype; subprolate.

Poilane 19777 [P] PI. V, 7 P =
32,5 n E 28,5 n P:E =
1,14.
Always two colpi transversales
per
Costae
colpus. 6-7 colpi. colpi.
Glochidion
type
Stephanocolporate (4); spheroidal -

prolate spheroidal.
Colpus transversalis circular or slightly elliptical; costae.
Colpi narrow.
lumina 1-2
Intectate
tectate; reticulate;
or
p.
In many morphological the Glochidion type is closely related

respects
to the Breynia type, as Erdtman (1952) and Webster (1956) have
stated. The lack, however, the double
pollen grains typical colpus
transversalis. It is also the in the Antidesma
possible to
place type
configuration. Its single circular, sometimes broad elliptic endexine
and the distinct reticulum are characters in favour of a

aperture
place in the Securinega type (p. 25).
Glochidion sericeum (Blume) J. U. Hooker Glochidion

type; spheroidal. P = E =
Bakhuizen v. d. Brink 1886 [U] 24 Colpi narrow. P.A.I. = 0,3-0,4.

n.
PI. V, 8 Lumina 1-2 /<• Intectate.
Glochidion concolor Muell. Arg. Glochidion type.

Yuncker 15648 [U]
Glochidion littorale Blume Glochidion type.
Bakhuizen v. d. Brink 3182 [U]
Glochidion obscurum (Willd.) Blume Glochidion type.
Buysman 295 [U]

Glochidion ramiflorum R. et G. Forster Glochidion
type.
Yuncker 15717 [U]
Glochidion Baillon Glochidion
superbum type.
Dumas 1507 [U]
40 W. PUNT
Glochidion spec. Glochidion type.

Teysman 4411 [U]
Arachnodes chevalieri Gagnep. Glochidion prolate spheroidal.
type;
Fleury 31755 PI. 1 P P:E
[P] VI, =
25,5 n
=
1,06.
P.A.I. > 0,5. Tectate; intra-reticu-
culate. Colpus transversalis slightly

elliptical.
Aristogeitonia configuration
Stephanocolpate, stephanocolporate, stephanoporate, periporate or inapertu-

rate. The pollen grains are spheroidal or have a Polar axis shorter than the
Equatorial axis.
Tectate; usually echinate, sometimes psilate.
The types of the

Aristogeitonia configuration form related
a
closely
which is differentiatedfrom the other in
group sharply configurations
the Phyllanthoideae. They the number of
are
easily recognised by
apertures and of the exine. Most have echinate
structure
types pollen
grains and ca. six small apertures. The shape is spheroidal or oblate
spheroidal to oblate.
The Aristogeitonia configuration includes a typical group

of genera
which in several different from other
are, respects, Phyllanthoideae.
Mueller
(1866) was acquainted with too few genera of this group
to recognise it. Bent ham (1880), however, enumerated, in the fourth
group of his Phyllantheae, all the genera of the Aristogeitonia configu-
ration then known. the
Only genus Bischofia of Bentham’s group
does the
not
belong to configuration.
All the the
genera belonging to
configuration are found in one of
the three into which Pax divides the
groups Phyllanthoideae (Pax 1924).
This includes his subtribes “Toxico-
group Drypetinae, Petalostigmatinae,
dendrinae”
(= Hyaenanchinae), Dissiliarinae, Paivaeusinae and the tribus
Caletieae of the Stenolobeae. and
Only Drypetes, Lingelsheimia Heywoodia
of the differ
Drypetinae pollen-morphologically. The genera are
small. The does than

largest genus Longetia not come up to more
10 species. The male flower lacks an extra staminal disc. The flowers
while the number of is
always are
apetal, stamens seldom constant;
there mostly than six. In the female flower the
are more
styles are
undivided, best but incised. The leaves of this

at emarginate, never
group are often opposite or

in whorls. Some genera have, moreover,
composite leaves.
Aristogeitonia type
Stephanocolpate, stephanocolporate or stephanoporate; oblate spherioidal -
suboblate.
Colpi, if present, small and

very short; costae colpi.
Pori, if with costae.
present,
Colpus transversalis, if
present, with costae.
Tectate; echinate or psilate.
In the Aristogeitonia type the apertures situated in the

are
always
pollen morphology of the euphorbiaceae 41
The have the of

equatorial plane. apertures can shape pori, colpi or
composite apertures, but they are always small.
Aristogeitonia subtype
Pollen grains echinate.
Aristogeitonia limoniifolia Prain Aristogeitonia subtype; stephanocolpate.

Herb. Kew. PI. VI, 5 P =
30,5 [i
E =
35 P:E =
0,86.
Colpi ca. 4,5 /i long. Echinae ca. 2,5 jx.
Oldfieldia africana Benth. Aristogeitonia subtype; stephanocolpate.

Small 621 [K] P -
35,5 E = 43 P:E =
0,83.
fi n
Paivaeusa dactylophylla Welw. et Oliver Aristogeitonia subtype; stephanocolpate.
Shabana 9 [K] P =
43
ft
E =
46,5 n
P:E =
0,92.
Piranhea trifoliata Baill. Aristogeitonia subtype; stephanoporate.
14238 PI. 4 P 27 E 32 P:E 0,85.
Glaziou [K] VI, =
fi
=
n
=
Diam. pori 3
fx.
Tetracoccus ilicifolius Coville et Gilmay
Aristogeitonia subtype; stephanocolporate.
Gilmay Mai. 2A. 1939 [L] P =
35,5 E = 40 P:E =
0,91.
fi
Colpi 6-7. Echinae 3,5 /x.
Hyaenanche globosa (Gaertn.) Lamb, et Aristogeitonia subtype; stephanocolporate.

Vahl P = 39 E = 46 P:E =
0,85.
ti n
Splitgerber [L] PI. VI, 6 Colpi 6-7. Echinae 2

ft.
Mischodon zeylanicus Thwait. Aristogeitonia subtype; stephanocolpate.
Hort IX A. 125. 80883 P 32 E 36 P:E 0,89.
Bogor. [U] =
n
=
n
=
Colpi 5-6. Echinae 3,5 fi.
Paragelonium perrieri Leandri Aristogeitonia subtype; stephanoporate.

Perrier de la Bathie 1178 [P] P =
32 n E = 35 P:E =
0,92.
v
Diam. pori 3 fi. Echinae 2,5-3,5 /x.
Petalostigma quadriloculare F. v. Mueller Aristogeitonia subtype; stephanoporate.

Queensland [U] P =
28,5 n
E = 32 « P:E = 0,89.
Pori 4-6; not always exactly in the
equatorial plane; diam. 3 Echinae

fi.
ca. 1 fi.
Longetia malayana (Benth.) Pax et Aristogeitonia subtype; stephanoporate.

K. Hoffmann P = 47
p
E =
52 ft
P:E = 0,90.
D’Alleizette, Penang 6421 [L] Pori slightly ellpitical diam. ca. 5
fi.
Echinae
up
to 6,5 fi.
Longetia carunculata (Baill.) Pax et Aristogeitonia subtype; stephanoporate.

K. Hoffmann Pori 5 (-6).
Mackee 4230 [L]
Longetia buxoides subtype
Pollen grains psilate.
Longetia buxoides Baillon Longetia buxoides subtype; stephanocolpate.

Nw. 6419 V 22 E=27 P:E 0,81.
D’Alleizette, Caled. [L] =
n n =
PI. VI, 7 Colpi 6 small and short

(5-) (-7);
with thick costae.
Dissiliaria tricornis Benth. longetia buxoides subtype; stephanoporate.

Brass 5755 [L] PI. VI, 9 P = 19 E =
22,5 P;E =
0,85.
/x. [x
Pori diam. 2
6; ft.
Stachystemon type
Periporate or inaperturate; spheroidal or ellipsoid (with a longer and a shorter
axis).
Pori small; costae pori.
echinate.
Tectate;
In the the situated in the

Stachystemon type apertures are never
equatorial plane, but scattered over the surface.

42 W. PUNT
plate vt. 1. Arachnodes chevalieri; 2. Phyllanthus nutans; 3. Dendrophyllanthus ripicolus;

4. Piranhea trifoliata; 5. Aristogeitonia limoniifolia; 6. Hyaenanche globosa; 7. Longetia
buxoides; 8. Phyllanthus acuminatus; 9. Dissiliaria tricornis.
vermicularis Planch. Stachystemon Diam. 36

Stachystemon type. pc.
Pritzel 270 [L] PI. VII, 2 Pori ca. 12; diam. pori 2,5 Echinae
pc.
1,5 jU long.
Micrantheum ericoides. Desf. Stachystemon type. Longer axis 45 fi. Pori
Buysman 1873; 347 [U] 25; diam. 2 Echinae in

ca.
fi. usually
of three around the pori; 3
groups n
long.
Micrantheum hexandrum D. Hook Stachystemon Diam. 45
J. type. pc.
Tasmania [U] Pori small; more than 40.
Pseudanthus pimeleoides Sieb. ex Spreng. Stachystemon Diam. 20

type. pc.
Buysman 1890 [U] PI. VII, 7 Pori ca. 7; diam. 2 pc.
Echinae 1 long.
pc
Pseudanthus nematophorus F. v. Muell. Stachystemon type. Diam 20 pc.
Murchinson [U]
banksii Benth. Stachystemon Diam. 40
Neoroepera type. pc.
Banks and Solander 1770 Diam. pori 1,5 30 pori.
[K] pc; ca.
Echinae 2 long.
pc
Longetia gynotricha Guile. Stachystemon type. Diam. 34 pc.

Mackee 4191 [L] Pori ca. 16. Echinae 2
pc.
Androstachys johnsonii Prain Stachystemon type.

Pole-Evans, 3-8-1917 [K] PI. VII, 2 Longer axis 46 Shorter axis 40 pc.
fi.
Echinae ca.
1
pc.
Pollen types not placed in one of the above configurations
Some cannot readily be placed in one of the discussed con-

types
figurations. Since most have only one
representative they
types
are discussed separately and not in any configuration.
put
Phyllanthus acuminatus type
Tricolporate (colpi diorate); spheroidal.

Colpi transversales (two in each colpus) circular; costae.
Colpi short.
Intectate; pilate. Pila ordinate arranged but not in a reticulum.
The presence of in the is also

two ora colpi an important character
The of the
of the
Breynia type (p. 38). structure exine, however, is
quite different. Its remaining characters cannot be compared with
any
other configuration.
Phyllanthus acuminatus Vahl Phyllanthus acuminatus

type.
Hostman 413 [U] PI. VI, 8 P =
E = 27 n. P.A.I. =
0,5.
Bischofia type
Tricolporate; spheroidal, P:E is 1.
Colpus transversalis small; costae. Edges of the costae transversales rounded
at the ends.
Colpi narrow and long; no costae colpi.

Tectate; psilate. Columellae short.
This type should be best placed in the Antidesma configuration but

differs from other in this its E
types configuration by shape (P: =
1)
and the absence of The columellae
costae
colpi. are moreover very
short.
44 W. PUNT
Bentham (1880) discussed with and

Bischofia together Oldfieldia
Piranhea because of their leaves. Pax
digitately compound and K.
Hoffmann (1931) thought it better to the in subtribe.

put genus apart a
As the similar those of

pollen grains are certainly not to Oldfieldia and
Piranhea and difficult in any other

moreover to place configuration,
it maintain the
seems right to separate subtribe.
Bischofia javanica Blume Bischofia type.

Hohenacker 1573 PI. 5 P E P:E 1,
[U] VII, = =
25 ju. = ca.
m:e < 0,5. P.A.I. =

0,2.
Putranjiva type
Tricolporate; spheroidal. P:E is 1.
Colpus transversalis large; costae.
Colpi narrow;
costae colpi.
Tectate; psilate. Exine thick. Columellae short but endexine and tectum thick.
The Putranjiva type has some resemblances with the Dicoelia

-
type
...
(p. 28). There is a large colpus transversalis with costae, and also
costae colpi. The type differs, however, by its shape (P: E

=1) and
the remarkably thick endexine.
In of
a
specimen Putranjiva (Stocks, India), examined by Erdtman
(1952), the pollen grains subprolate, reticulate. The
are
specimens
Reorders 2156 and from that
Broadway 9249 differ considerably
description. The typical character of the thick exine is not mentioned
Erdtman and in his

by was
probably not
present pollen grains.
In the system of Pax and K. Hoffmann (1931) Putranjiva is found

in the subtribe Glochidiinae. The is in
genus put relatiosnhip with
and Glochidion.
Breynia Bentham (1880) gives it a
place next to
It be admitted, that the of

Drypetes. must pollen grains Putranjiva are
far similar those of than the

more to
Drypetes to
Breynia type.
Putranjiva roxburghii Wall. Putranjiva type. P == E = 40

fi.
Koorders 2156/3 [U] P:E = ca. 1. m:e<0,5.
Braodway 9249 [U] PI. VII, 3 P.A.I. 0,3. Exine thick
=
(4 fi).
Hymenocardia type
Triporate (-diporate); oblate.
Costae pori.
The type differs from all the other in the

completely pollen grains
Phyllanthoideae.
On account of its catkin-like male inflorescenses Pax and K.
Hoffmann
(1931) placed Hymenocardia in the Antidesminae. The
character of the is the

most striking genus compressed two-cocced
fruit with lateral wings. Hymenocardia differs from the other genera in
the Antidesminae in some
characters:
I. Androstachysjohnsonii; 2. Stachystemon vermicularis; 3.

plate vh.
Putranjiva roxburghii;
4. Martretia quadricornis; 5. 6. 7. Pseudanthus
Bischofia javanica; Hymenocardia ulmoides;
pimeleoides.
46 w. PUNT
1. Disc in male well in female flowers.

wanting as as
2. The styles are long and undivided.
Hymenocardia ulmoides Oliv. Hymenocardia type.

Koechlin 2371 [U] PI. VII, 6 P = 19 E =
22,5 P:E =
0,85.
fi n
Hymenocardia acida Tul. Hymenocardia type.

Bequaert 7142 (BR) P = 28,5 /j E = 31 P:E =
0,92.
fi
Duvigneaud 2422 (BRLU) Diam. pori 3

fi.
Martretia type
Tricolporate; oblate
spheroidal.
Colpus transversalis; costae. Edges of the costae transversales rounded at the
ends. Atrium present.
Colpi narrow; no costae colpi present.

Tectate; psilate.
The of the Martretia oblate but

pollen grains type are spheroidal
do not have any other character in common with the Amanoa or
The has rounded ends.

Aristogeitonia configuration. colpus transversalis
This character is present in some
types of the Antidesma configuration.
and absence of
The oblate shape costae colpi keeps the Martretia type
out of the Antidesma configuration. A typical character is the atrium,
which is not found in any other
type.
Martretia has only one species. In many respects this species is

related to the Antidesminae (e.g. in its inflorescense). There are, however,
some differences.
1. No disc is present in the male flower nor in the female one.
2. The seeds are provided with a caruncula. No other in the

genus
Antidesminae has carunculate seeds.
3. The in the whole

only genus family with a false partition in the
fruits (Pax and K. Hoffmann, 1922).
Martretia quadricornis Beille Martretia

type.
Adames 251 [K] PI. VII, 4 P 28,5 E 31 P;E
=
n
=
n
=
0,92.
m:e = 0,3 —
0,4. P.A.I. = 0,25.
Taxonomic comment on the

Phyllanthoideae
The Phyllanthoideae can be divided into three groups of pollen types.

These broadly outlined, with the three groups Pax
groups, agree
(1924) could already distinguish on other grounds, though without
nominating them.
The first for the subtribes Antidesmineae
group, to which, instance,
and has which have
Phyllanthinae belong, pollen grains mostly a
and distinct transversales

prolate shape costae ( Antidesma configuration,
p. 20). The second group has pollen grains which are mostly reticulate
and generally have oblate shape (,Amanoa configuration,
an p. 32).
The subtribes Amanoinae, Sarropodinae, etc., belong to this group.
The third comprises those genera
that are to be found in the
group
Aristogeitonia configuration (p. 40).
For the
Phyllanthoideae (Pax and K. Hoffmann, 1931) composed a
highly differentiated system of many

subtribes. This system, however,
leaves much to be desired. Some subtribes (e.g. Glochidiinae) comprise
which do show with each
genera decidedly not
any relationship
other. On the other subtribes resemble
contrary, genera in different
each other so much, that it would seem desirable to unite them
(Wielandiinae with Amanoinae and Pseudolachnostylidinae).

In the
Phyllanthoideae pollen morphology is obviously exceedingly
useful to arrive at a natural classification.
b. Crotonoideae
Croton configuration
Croton-pattern present. Structure elements on the tectum are usually clavae
but can also be echinae, baculae etc. Sometimes the structure elements are
located ridges (e.g. Croton matourensis, Manihot Tectate.

on
saxicola).
The Croton includes all

configuration pollen grains with a croton-
In the the is found in the

pattern. Euphorbiaceae croton-pattern only
the Crotonoideae. Erdtman outside the
subfamily of (1952) states, that,
Euphorbiaceae, pollen grains with a have been found
croton-pattern
in the Buxaceae and
Thymelaeaceae.
Most
pollen grains with a
croton-pattern belong to the Croton
type.
The Manihot type, Klaineanthus and Hevea occur
less
commonly.
type type
In the subfamily of the Crotonoideae a great number of genera possess

pollen grains with a croton-pattern.
In Pax and K. Hoffmann’s the
system (1931) these genera occur in
following tribes: Grotoneae, Ghrozophoreae, Joannesieae, Cluytieae,

Manihoteae, Celonieae and Ricinocarpeae. Further also Neoboutonia
as the only genus out of the

Acalypheae. It follows from this enumera-
tion, that the genera possessing a croton-pattern are to be found in
widely divergentgroupings.
In authors considered the
classifying the Crotonoideae most bursting
of the male be factor of primary for
open calyx to a
importance
arranging the genera. Thus Mueller (1866) and Pax and K.Hoff-
mann arrived at a classification, in which with an obvious

genera
relationship are kept widely Joannesia and Micrandra

apart, e.g.
Schultes Schultes
(Baldwin et 1947; 1955). Bentham (1880)
already stated that within the genus
Croton the male calyx opens
both
valvate and which is in his
imbricate, why, system, this character
less In Bentham’s the genera with

plays a
important part. system a
croton-pattern are placed close together. They are to be found in his
subtribes Jatrophineae, Eucrotoninae and Ghrozophorinae.

In all the that it is
comparing taxa
possess a croton-pattern seen
that several characters correlate with this character. With

pollen
hardly an
exception the male and the female flower contain a disc,
while ovarium rudiment is absent. A less but
an constant striking
48 W. PUNT
character is the of
presence petals in many genera.
Genera which
form an exception to these characters are:
disc is in both
Sagotia. The wanting sexes. The genus, however, is
related Sandwithia and Garcia
to Dodecastigma, (Croizat 1948), which,
on the other hand, have a
disc.
has disc in both

Bertya. This Australian genus no sexes either. There
distinct relation other Australian genera, viz.

is, however, a to two
and which have disc.

Beyeria Ricinocarpos, certainly a
Tritaxis has no disc in the female flower.
Adenocline and Tetrorchidium have no disc in the male flower.
The Pantadenia and described

genera Oligoceras, by Gagnepain, were
said to
possess an ovarium rudiment in their male flowers. Pantadenia
does have little conical in the of its
a projection centre receptaculum,
but whether this ovarium rudiment is
projection represents an not
certain. in its male column

Oligoceras possesses, flower, a which,
Gagnepain in the Flore General de
according to a
drawing by
ITndochine (1926), was said to be an ovarium rudiment. In studying
the material in Paris, however, the author concluded, that the
type
the The column is
drawing gives an erroneous picture of reality. not
but The five

bottle-shaped cylindrical. stamens are not placed at
the foot of the column, but are grown together with it at a little
distance of the base. The

projections which stand at the end of the
column and which, in the drawing, look like three

stigmas, are
staminodies. These staminodies even contain rudimentary pollen sacs.
Therefore there are two whorls of stamens, the topmost of which is
sterile.
A more or less distinct ovarium rudiment is found in:

all
Klaineanthus, Cladogelonium, Micrandra and Endospermum, which belong
to the Klaineanthus type.
Genera not possessing a corolla are: Neoboutonia, Benoistia, Elaterio-
spermum, Baliospermum, Eremocarpus, Micrandra ( Cunuria ), Hevea, Bertya,

Beyeria, Adenocline and Tetrorchidium. Suregada mostly has no corolla
(Leonard 1958).
The is the Croton Less
type occuring most frequently type. common
the Manihot the Klaineanthus Hevea Adenocline

are
type, type, type and
type.
Croton type
Inaperturate; spheroidal or ellipsoid i.e. with a longer and a shorter axis.

Tectate; endexine thin. Croton-pattern.
Clavate or
echinate.
The Croton has and the

type no
apertures shape is fairly uniform.
There may be a longer and a shorter axis, but usually the pollen grains
are spheroidal. The structure elements have more variations. The
differences in these elements and the lumina enclosed
by the elements,
however, are slight and many
transitions exist, so that it seems better
omit the formation of groups
to (subtypes).
of
The
pollen grains Sagotia racemosa show two
shapes. The specimens
Y. Mexia and Woob Herb, are echinate, while the specimen For. Dep.
1. Croton
plate vm. cuneatus; 2. Croton matourensis (x 800).
50 w. PUNT
is clavate. Further examination will be to ascertain whether

necessary
these of be other
two
shapes Sagotia racemosa can distinguished by
characters besides.
Schultes be from
According to
(1952) Cunuria cannot distinguished
Micrandra. The pollen grains of the Cunuria specimen Ducke 1087
the Croton In another

belong to type, and not to the Micrandra
type.
Cunuria specimen (Ducke 24874) Erdtman
(1952) described the pollen
Schultes the
grains as being 5-colporate. According to (1952) two
Ducke specimens both belong to Cunuria var. bracteosa.

spruceana
Micrandra and Micrandra stand
siphonoides brownsbergensis pollen-
morphologically between the two specimens of Cunuria spruceana var.
bracteosa.
Klotzsch Croton
Croton cuneatus type.
PI.
J. et P. Florschutz 1113 [U] VIII, 1 Diam. longer axis 75
/*. Clavate; diam.
clavae 3,5 fi.

Clavae on ridges.
Millsp. Croton Diam. 55
Croton bahamensis type. /x.
Brace 4064 [U] D. cl. 2,8 fi.

Croton longiradiatus Lanj. Croton
type.
Diam. 75 fi.
B.W. 6711 [U] D. cl. 2,2 Clavae distinctly on ridges.

/x.
Croton hirtus L’Heritier Croton type.
504 [U]
Versteeg
Croton matourensis Aubl. Croton
type.
B. 5050 [U] PI. VIII, 2 Clavae acuminate; distinctly on ridges.
Diam. 115 D. cl. 6,5
fi. fx.
Croton pullei Lanj. Croton type.
Rombouts 654 [U] Diam. 115 D. cl. 6,5 Clavae
fi.
distinctly on
ridges.
Julocroton triqueter Baillon Croton type. Diam. 120
/x.
6016 [U] D. cl. 3,2
Hassler fx.
Julocroton argenteus (Linn.) Didrichs Croton type. Diam. 120 /x.
C. L. Schulz 1225 [U] D. cl. 3,2 fx.
Crotonopsis elliptica WlIXD. Croton type. Diam. 60

fx.
Delzie 7066 [U] D. cl. 1,5
Demaree fx.
Benth. Croton type. Diam. 60
Eremocarpus setigerus (Hook.) /x.
L. S. Rose 41457 [U] D. cl. 2 /x.

Pax Croton Diam. 48
Grossera major type. fx.
Zenker 469 [U] D. cl. 1,8 /x.
Holstia tenuifolia Pax Croton type. Diam. 45

fx.
Hemsley 3480 [K] D. cl. 2

Drummond et fx.
Cyrtogonone (Pax) Prain Croton Diam. 60

argentea type. /x.
Zenker 561 [U] D. cl. 3,5 fx.
Crotonogyne preussii Pax Croton type.

Diam. 60 /x.
Zenker 4666 [L] D. cl. 2

_ /x.
Crotonogyne parvifolia Prain Croton Diam. 50

type. fx.
D’AlTeizette 6845 [L] D. cl. 2,8 /(.
Manniophyton fulvum Muell. Arg. Croton type. Diam. 60

/t.
Zenker 1404 [L] D. cl. 3,5 /x.
Aleurites moluccana (Linn.) Willd. Croton type. Diam. 70

fx.
Bunnesmeyer 8060 [U]

D. cl. 2,0 /x.
Deutzianthus tonkinensis Gagnep. Croton type. Diam. 65

/x.
Balansa 3149 [P] D. cl. 2,0 /x.
Gagnep. Croton Diam. 50
Oligoceras eberhardtii type. /(.
Eberhardt D. cl. 1,8

[P] fx.
Alphandia furfuracea Baillon Croton

type. Diam. 40 /x.
Balansa 3439 [P] D. cl. 2,0 /.i.
Garcia nutans Vahl Croton Diam. 60

type. /(.
Lundell 12248 [U] D. cl. 2,0 fx.

Velloso Croton Diam. 70

Joannesia princeps type. ft.
T. W. Brown 35 D. cl. 3,0 Clavae ridges.

[K] ft.
on
Pax Diam. 40
Neoboutonia macrocalyx Croton type. ft.
Stolz 2043 D. cl. 1,5

[U] fi.
Pantadenia adenanthera Gagnep. Croton Diam. 45

type. ft.
British Museum D. cl. 1,0 fi.
lucida Endlicher Croton Diam. 40

Baloghia type. fi.
Hastings River [U] D. cl. 1,0

Beckler, ft.
Blume Croton Diam. 65

Ostodes paniculata type. ft.
Griffith 4790 [U] D. cl. 3,5 ft.
Meeuse et Croton Diam. 53

Ostodes pendulus (Hassk.) type. ft.
A DIPT RFRT D. cl. 2,8 ft.
Sandakan For. Dep. A. 2964 [U]

Codiaeum Blume Croton Diam. 50
variegatum (Linn.) type. ft.
Fresh material from greenhouse at D. cl. 2 ft.
Utrecht. PI. IX, 3
Codiaeum Ware. Diam. 50

stellingianum Croton type. ft.
Docters v. Leeuwen 1567 _[U] D. cl. 2,0 ft.
Heckel Croton Diam. 60

Fontainea pancheri (Baillon) type. ft.
D’Alleizette 6474 [L] D. cl. 2,8 ft.
murina Elmer Croton Diam. 80

Dimorphocalyx type. ft.
Elmer D. cl. 3,5

12773 [U] ft.
Benoistia Leandri Croton Diam. 35

perrieri type. ft.
D’Alleizette Dec. 1905 [L] D. cl. 1,8

Madag. ft.
Diam. 45
Tritaxis gaudichaudi Baill. Croton type. ft.
Clemens 4351 Echinate. Diam. ech. 1,8

[U] ft.
Boerl. Croton Diam. 60

Strophioblachia fimbricalyx type. ft.
Sort. VIS F. 43 [U] D. cl. 1,8

Bog. ft.
Baill. Croton Diam. 45

Blachia umbellata (Willd.) type. ft.
D’Alleizette Austr. Nov. 1907 [I.] D. cl. 1,8 ft.
Baillon Croton Diam. 50

Sagotia racemosa
type. ft.
For. Dep. 7443 [U]

D. cl. 2,0. Clavae sharp pointed.
Mexia 6050 Echinate. Diam. 40 D. ech. 1,8

[U] ft. ft.
Wood Herb. 311 [U]

Linn. Diam. 95
Jatropha multifida Croton type. ft.
St. Eustatius [U] D. cl. 4,0 Clavae ridges.

Groll-Meyer, ft. on
Croton Idem. Jatropha multifida

Jatropha integerrima Jacq,. type.
Docters v. Leeuwen 19-7-1950 [U]
Mildbraedia Pax Croton Diam. 50
paniculata type. ft.
Leeuwenberg 3004 [U] D. cl. 2,5 ft.

Crises. Croton Diam. 45
Acidocroton adelioides type. ft.
Ekman 16896 D. cl. 1,5

[K] ft.
Pausandra densiflora Lanj. Croton Diam. 50

type. ft.
Krukoff 5464 D. cl. 2,0

[U] ft.
Pausandra morisiana Radlk. Croton Diam. 75

(Casar.) type. ft.
Ducke et Kuhlmann 16594 [U] D. cl. 3,5 ft.
Griffith Croton Diam. 48

Givotia rottleriformis type. ft.
2638 D. cl. 2,8

Wight [L] ft,
Ricinodendron heudelotii (Baill.) Pierre Croton Diam. 50

type. ft.
D. cl. 2,5 ft.

Zenker 487 [U]
Blume Croton Diam. 70
Elateriospermum tapos type. ft.
cl. 3,2 Clavae
Hort. Bog. IX. A. 129 [U] D. ft.
on
ridges.
MUELL. Croton Diam. 43
Baliospermum motanum (WlLLD.) type. ft.
D. cl. 1,5 ft.
D. Hooker [U]
Bengal, J.
Leonard Croton Diam. 75
Cavacoa aurea (Cavaco) type. ft.
de Winter 2109b [K] D. cl. 2,0 ft.
uleanus et K. Hoffm.) Croton Diam. 55

Anomalocalyx (Pax type. ft.
OlTPIfP D. cl. 3,2 ft.
Ducke 23518 [U]

52 W. PUNT
plate rx. 1. Manihot saxicola (X 600); 2. Tannodia cordifolia; 3. Codiaeum variegatum.

POLLEN MORPHOLOGY OF THE EUPHORBLAGEAE 53
Sandwithia guyanensis Lanj. Croton Diam. 55

type. fi.
For. Dep. 2731 [U] D. cl. 3,5 fi.
Dodecastigma amazonicum Ducke Croton Diam. 80

type. ft.
Ducke 23543 [U] D. cl. 4,5 ft.
Cunuria
spruceana Baillon var. bracteosa Croton type.
Diam. 50 fi.
(Ducke) R. E. Schultes D. cl. 1,8 fi.

Ducke 1087 [K]
Vaupesia cataractarum R. E. Schultes Croton Diam. 70
type. fi.
Schultes 14006 D. cl.
[K] 3,0 ft.
Ricinocarpos pinifolius Desf. Croton

type.
Diam. 60 fi.
Austr. felix [U] D. cl. 1,8 ft.
Beyeria leschenaultii (Decand.) Baill. Croton type.

Diam. 43
fi.
Austr. felix [U] D. cl. 1,5 u.
Clavae sharp pointed.

Bertya gummifera Planch. Croton Diam. 55
type. fi.
Johnson et Constable 16024 Croton-pattern indistinct.

[U]
Domohinea perrieri L£andri Croton Diam. 50
type. ft.
D’Alleizette 6479 Dec. 1905 [L] PI. X, 1 Croton-pattern on ridges. Echinate;
diam. ech. 1,5 fi.
Tannodia cordifolia Baillon Croton Diam. 35
type. fi.
D’Alleizette, Comoren 6484 Echinate; diam. ech. 1-1,5

[L] fi.
PI. IX, 2
Manihot
type
Periporate; spheroidal.
Pori large, borders indistinct.
Tectate. Croton-pattern.
The type is related the Croton type. The

closely to
pollen grains,
however, have large with indistinct rims. The number of
apertures
varies in the different
pori species.
According Croizat (1943b) Cnidiscolus does have

to not a calyx
but a corolla. This is also the case with Manihot. In a definite
very
sense Cnidiscolus connects Jatropha, sensu strictu, with Manihot. Both
the of Cnidiscolus examined had of the Manihot

sorts
pollen grains
type, sothat least in this respect closer related Manihot
at
they are to
than to Jatropha.
The
pollen-morphological similarity between Manihot and Suregada
is probably accidental and has to be considered a parallel development.
Manihot saxicola Lanj. Manihot

type.
Proefst. Buitenz. 922 Boom 8 Diam. 160 Croton-pattern
[U] fi. on a
PI. IX, 1 distinct reticulum.
Pori ca. 25. Diam. clavae 6,8 fi.

Manihot esculenta Crantz Manihot
type.
Rombouts 757 [U] Idem Manihot saxicola.
B.W. 2780 Pori 25.

[U] ca.
Cnidiscolus Arthur Manihot Diam. 75

urens
(Linn.) type. fi.
Boldingh 4829 [U] D. cl. 2,5 fi.

Pori 3^1.
Cnidiscolus stimulosus Gray Manihot Pori 6.

(Michaux) type. ca.
Killip 31636 [U]

Suregada glomerulata (Blume) Baill. Manihot Diam. 40
type. ft.
Griffith 4772 [U] D. cl. 2,5 Pori 3-5.
fi.
Suregada subglomerulata (Elmer) Croizat Manihot

type. Diam. 45
/i.
Elmer 12967 [U] Pori 5-6.
54 W. PUNT
Klaineanthus type
Tricolpate; oblate spheroidal to oblate.
Colpi wide or no operculum present.

narrow;
Tectate; croton-pattern.
The Klaineanthus type, related the Croton

too, is closely to type and
the Manihot Pollen grains of this type have colpi without an
type.
The membrane is
operculum. colpus frequently ruptured.
Klaineanthus
subtype
Boarders of colpi distinct.
Glavate.
The rims of the of this distinct. The

colpi subtype are colpus
membrane is frequently totally ruptured. In Cladogelonium the
colpus
membrane is but Sometimes rudiment of
not ruptured granulate. a
the colpus membrane remains at the outer ends of the colpi.
Klaineanthus gaboniae Pierre ex Prain Klaineanthus subtype; suboblate.
Zenker 583 [U] PL X, 3 P =

29,5 £ =
37/1 P:E = 0,80.
/i
Colpi wide. P.A.I. =

0,4.
Diam. clavae 1,8 /t.
madagascariense L6andri Klaineanthus subtype; spheroidal.

Cladogelonium
Perrier de la Bathie 9696 [P] PI. X, 4 P =
30/i E =
30,5 P:E =
0,98.
fx
Colpus membrane granulate.

P.A.I. =
0,4.
Endospermum moluccanum (Thysm. et Klaineanthus subtype; suboblate.
Binnend.) Beccari P =
33,5 E =
44,5 P:E =
0,80.
n /i
Kornasse 462 [U] Colpus membrane absent. P.A.I. =
0,4. Diam. clavae 1,8 /i.
Glycydendron amazonicum Ducke Klaineanthus subtype; oblate.
Ducke 17108 [U] P =

29 E =41,5/t P:E =
0,69.
/«
Colpus membrane absent. P.A.I. =
O, Diam. clavae 1,5 fi.

Micrandra brownsbergensis Lanj. Klaineanthus subtype.
v. Emden 9 [U] Colpus membrane absent.
Micrandra siphonioides Benth. Klaineanthus subtype.
Lopez 9625 [U] PI. X, 5 P =

41,5 fi E = 46 /< P:E =
0,92.
P. =
0,3. Croton-pattern indis-
tinct. Clavae candelabrum-shaped,

in-ordinate arranged.
Adenocline
subtype
Borders of the colpi not sharply cut but irrigularly ruptured.
Baculate. Baculae small and crowded.
The baculae on the tectum are

small and crowded. With low
magnification it is obvious that the elements

not
directly structure
form a and not a reticulum.

croton-pattern
The Adenocline
genus was placed in the vicinity of Seidelia and
Leidesia in the Acalypheae by Bentham (1880). Pax and K. Hoffmann
(1931) could not find the same relation and put the genus in a
separate
subtribe in the tribe Gelonieae. This tribe also the
comprises genus
Klaineanthus. The two

genera are, however, quite different in other
respects.
POLLEN MORPHOLOGY OF THe'eUPHORBIACEAE 55
plate x. 1. Domohinea perrieri; 2. Suregada subglomerulata; 3. Klaineanthus gaboniae;

4. Cladogelonium madagascariense; 5. Micrandra siphonioides; 6. Hevea brasiliensis.
56 W. PUNT
Adenocline acuta Baiix. Adenocline subtype;

(Thunb.) oblate spheroidal.
[= A. mercurialis Turcz.] P 36 E 37,5 P:E
=
/a =
n =
0,96.
Lanjouw 613 [U] PI. XI, 2 P.A.I. = 0,4.
Hevea type
Tricolpate; oblate spheroidal.

Colpi wide; operculum present.
Tectate; baculate. The baculae are so small and crowded, that the structure
is hardly recognised as a
croton-pattern.
As in the Adenocline subtype the baculae of the Hevea type are small
in diameter and crowded. With low magnification the structure ele-

form reticulum, but with
ments seems to a
high magnification the
of the structure elements is clearly a
arrangement croton-pattern.
The operculum distinguishes this type from the Klaineanthus
type.
Hevea brasiliensis (Willd. Hevea

ex A. Jussieu) type; oblate spheroidal.
Mueix. Arc. P 33,5 E 35 P:E
=
fi
=
n = 0,96.
Schultes 8050 [U] P.A.I. = 0,3.
B.W. 752 [U] PI. X, 6
Hevea guianensis Aublet Hevea

type.
B.W. 41 fU] P =
35 E = 38 P:E =
0,95.
i* n
Krukoff 5800 [U]
Tetrorchidium
type
Tricolporate; oblate spheroidal to prolate spheroidal.

Colpus transversalis exceptionally broad and elongated.
No costae present.
Colpi rims irregular.
narrow;
Intectate; pilate. Pila coarse.
The Tetrorchidium is and differs

type tricolporate by this character
from the Klaineanthus The
tricolpate type. extremely broad colpus
transversalis is elongated transversalis The
to a
colpus equatorialis.
tectum of this is indistinct and the is clear
type croton-pattern not
in all the In Tetrorchidium the

pollen grains. didymostemon structure
elements on the tectum are certainly not arranged in a

croton-pattern.
Tetrorchidium rubrivenium shows the croton-pattern more readily.
Although the important characters of the Croton configuration

most
are not always present, the structure of the pollen grains and the shape
of the much like those in the Klaineanthus
colpi are very type. The
_
author therefore the Tetrorchidium type in the

prefers to
keep vicinity
of the Klaineanthus type.
Pax and K. Hoffmann referred Tetrorchidium into

(1931)
v ,
the sub-
tribe Tetrorchidiinae. This subtribe was placed in the tribe Gelonieae,

to which also Klaineanthus belongs. Although in habit and character
Adenocline and Tetrorchidium differ the flowers have
widely, some
characters in common:
1. In the male flowers disc and ovarium rudiment are absent.
2. In the female flowers the disc consists of three free glands.

Tetrorchidium rubrivenium Poepp. Tetrorchidium oblate spheroidal.

type;
Harris and Britton 10746 P 36 £
[K] =
n
=
40/1 P:E =
0,90.
P.A.I. = 0,35. Bacculate-verruca te.
Tetrorchidium didymostemon Pax Tetrorchidium

(Baillon) type; prolate spheroidal.
et K. Hoffmann P = 28 /i E =
27 n P:E =
1,04.
Zenker 4590 [L] PI. XI, 3 P.A.I. =
0,35.
Cnesmosa configuration
Inaperturate; no croton-pattern present.

Tectate or intectate.
Like the Croton the Cnesmosa configuration is inaperturate, but

type
a is always absent.
croton-pattern
Cnesmosa type
Pollen grains ellipsoid, i.e. with shorter

Inaperturate. a longer and a axis,
Tectate; psilate. No reticulum.
The of this
pollen grains type
are
closely related to the Tragia fallax
and the Platygyne type. In the Cnesmosa the
type type pollen grains
are tectate.
The genera belonging to this type are

closely related and not
easy
to separate. Croizat (1941a) discussed the relationship between some
genera of the Plukenetiinae and came to the following conclusions:
1. Cenesmon Gagnepain be from Cnesmosa Blume.

cannot
separated
2. Megistostigma malaccense, reduced to Sphaerostylis by Pax and
K. Hoffmann, is the taxonomic

type of the genus Megistostigma,
which is congeneric with Clavistylus J. J. Smith.
3. Tragiella Pax and K. Hoffmann should be treated as a synonym

of Sphaerostylis, of which S. tulasneana is the taxonomic type
(p. 63).
The results in full with the

pollen-morphological are
agreement
statements of Croizat.
Cnesmosa javanica Blume Cnesmosa ellipsoid.

type;
Elmer, Borneo 20663 [U] Longer axis 55
p.
Java 1855 [L]

Megistostigmapeltatus (J.J. Smith) Croizat Cnesmosa
i ellipsoid.
type;
Steenis 1235 [L] PI. 6 axis 56 shorter axis 53
v. XI, Longer fi ; p.
Megistostigma malaccense J. D. Hook. Cnesmosa ellipsoid.

type;
Sinclair 40242 [U] PI. XI, 4 Longer axis 42 shorter axis 37
//; /i.
Columellae in
groups.
Acidoton Swartz Cnesmosa ellipsoid.
urens
type;
Jamaica, Hooker axis shorter axis
[U] Longer 37,5 fi \ 35,5 ft.
fallax
Tragia type
Pollen grains inaperturate; spheroidal ellipsoid. Pila

or Intectate, pilate. not
in a reticulum. Endexine thin.

very
This type differs from the Cnesmosa type by its intectate structure.
The in reticulum.
pila are not arranged a
58 W. PUNT
The examined the section Bia of Tragia. Klotzsch

species belong to
and Baillon this section from and K.

(1858) separated Tragia. Pax
Hoffmann (1919a) also stated: “Die mannliche Bliite weckt den
als ob eine
Eindruck, es um eigene Gattung handelt’Mf all the
species
of the section Bia have the it is
inaperturate pollen grains, perhaps
better this section distinct genus,
to treat as a although a
great affinity
with Tragia cannot be denied.
Tragia fallax Muell. Arc. Tragia fallax ‘ype

-
;
Klug 4207 [U] PI. XI, 5 Diam. 50 u. Pila 1,5 /J.
Tragia sellowiana (Klotzsch) Muell. Tragia fallax type.

Arc.
Smith 3558 [U] Diam. 55

/a.
Platygyne type
Inaperturate. Pollen grains ellipsoid.

Tectate; intra-reticulate. Endexine with circular thickenings.
The is distinguished from the Cnesmosa its

Platygyne type type by
distinct intra-reticulum. Small circular thickenings occur on the
endexine. These peculiar “costae” are distributed irregularly over the
endexine and certainly not accompagnied by thinnings of the ektexine.
Platygyne hexandra (Jacq.) Muell. Arg. Platygyne ellipsoid.

type;
Rutten-Pekelharing 346 PI. 1 axis 40 Shorter axis 36
[U] XI, Longer fx. u.
Lumina 2-3 fi.
Trigonostemon verrucosus type
Inaperturate; spheroidal or ellipsoid.

Tectate. Gemmate.
Trigonostemon verrucosus J. J. Smith Trigonostemon verrucosus

type; spheroidal.
Longer axis 60 Diam. 2
/(. gemmae jx.
Cult. Hort. Bog. VIII. E-16 [U] Gemmae not in a

croton-pattern.
PI. XII, 3 Trigonostemon verrucosus
type.
Trigonostemon fungii Merrill Diam. 60 Diam. 1,5-2
fi. gemmae /i.
Linsley Grishitt 1089 [BM]
redioides
Trigonostemon type
Inaperturate; spheroidal to elliptical.

Intectate; reticulate.
Both
Trigonostemon types are inaperturate without a
croton-pattern.
They belong for these reasons to the Cnesmosa configuration.
The with the
Trigonostemon verrucosa
type gemmate structure
elements its is related the Croton

on tectum
perhaps to
type (p. 48).
The
gemmae are, however, certainly in
not
arranged a
croton-pattern.
the and lack of the
Except for shape apertures Trigonostemon redioides
does not have character in common with the Croton con-
type any
figuration.
plate xi. 1. Platygyne hexandra; 2. Adenocline mercurialis; 3. Tetrorchidium didymostemon;

4. Megistostigma malaccense; 5. Tragia fallax; 6. Megistostigma peltatus.
60 w. PUNT
the of the
Although pollen grains _
Trigonostemon type
_
are not provided
—
with a
croton-pattern, the genus Trigonostemon has great affinity with
in the Croton characters
genera configuration. Corresponding are:
1. In the male as well in the female flowers a disc is found and an
ovarium rudiment is absent.
2. Sepals are
present.
Trigonostemon redioides (Kurz) Craib Trigonostemon redioides Longer axis

type.
Coudre [BM] PI. XII, 5 34 Shorter axis 30 Lumina 1
fi. ft. -2 ft.
Trigonostemon longifolius Wall, ex Baillon Trigonostemon redioides type-

Achmad 142 [U] Diam. 55 Lumina 3-5
ft. fi.
Prosartema gaudichaudii Gagnep. Trigonostemon redioides

type.
Poilane 10446 [K] Longer axis 38 snorter axis 35
p\ fi.
Lumina ca. 2 fi.
Actephilopsis malayana Ridl. Trigonostemon redioides

type-
Ridley 2300 [K] Longer axis 45 Lumina small,
fi.
ca. 1
/i.
Dysopsis configuration
Inaperturate. Pollen grains tricolpate, distinctly three-lobed.

quasi
Tectate; no
croton-pattern present.
Dysopsis type
Inaperturate; quasi tricolpate; oblate spheroidal.

Tectate; psilate. Columellae fairly coarse.
The striking character is found in the three-lobed of

most
shape
the The declination between the lobes is but
pollen grains. distinct,
there is the indication of any
not slightest part of the exine being
weakened. the definition the
According to (see glossary) pollen grains
are inaperturate.
Mueller well
(1866) as as Pax and K. Hoffmann
(1914, 1931)
in close with Leidesia and Seidelia. These
kept Dysopsis relationship
latter African however,
genera are, geographically far removed from
the Andine genus The three genera

Dysopsis. belong undoubtedly to
the The have

Acalypheae. pollen grains of Dysopsis little affinity with
those of Leidesia and Seidelia. it should be better
Perhaps to
separate
in different subtribe.
Dysopsis a
Dysopsis glechomoides (Rich.) Muell. Arg. Dysopsis type.

.
. „ .
. ,
Lechler 956 [K] PI. XII, 6 P 21 E 22,5 P:E 0,93.

/i fi
= =
=
Plukenetia configuration
Tricolpate triporate. Shape of the pollen grains oblate oblate

or
spheroidal to
(rarely prolate spheroidal).

Apertures broad; costae absent.
Colpus or
porus
membrane ruptured.
1. Plukenetia 2. Fragariopsis scandens; 3.

plate xn.
verrucosa; Trigonostemon verrucosus;
4. Plukenetia volubilis; 5. Trigonostemon redioides; 6. Dysopsis glechomoides.
62 W. PUNT
The of the Plukenetia have

types configuration simple apertures.
In with the the
contrast Chiropetalum configuration apertures are
broad and have In

a ruptured colpus membrane. some species a small
of this membrane the ends of the
piece persists at outer colpi.
Plukenetia
type
Tricolpate; oblate spheroidal to oblate (rarely spheroidal or prolate spheroidal).

Colpi broad; colpus membrane ruptured. At the outer ends of the
apertures
small piece of the colpus membrane
a
(i.e. endexine) persists.
The is sometimes bordered
ruptured part by a
margo.
Tectate or intectate. Reticulate or not reticulate.
Pollen this have different exine

grains belonging to type can very
be while
structures. They may intectate, tectate or tectum perforate,
the columellae inordinate form reticulum. The
are
arranged or a
various subtypes are based on the structure patterns of the exine.
From the Plukenetia new

have been separated by
genus genera
j
several authors. 1 ax and K. Hoffmann maintain this division
~ (1919a)
of Plukenetia. Bentham
(1880), however, has pointed out that Fragariop-
sis and Romanoa differ only slightly from the typical Plukenetia species.
Other authors, too, see little difference. Uroizat (1941a): “JNo
characters are available to Pterococcus and Tetracarpidium

separate
“
from Plukenetia Macbride and
(1951): Apodandra Fragariopsis
could be included in Plukenetia". the
readily Certainly pollen grains
of the above genera are not
easily distinguished from those of Plu-
kenetia.
In South-America three
tree-shaped, non-climbing genera are to
“
be found. Sandwith of them: Astrococcus and
(1950) says Angostyles,
Haematostemon three allied genera of the Plukenetiinae”.
are closely
The all
pollen is indeed related. They belong to the Plukenetia volubilis
and and Haematoste-

subtype possess a tectum perforatum. Astrococcus
mon have moreover a
margo.
On of the imbricate Bentham and Pax and K.
account calyx
Hoffmann
placed Omphalea outside the Plukenetiinae. Croizat (1942a,
however, his that the shows
1948), gives as opinion genus strong
affinity to the Plukenetiinae. The pollen of Omphalea also appears to
resemble the Plukenetia volubilis subtype very closely.
Plukenetia volubilis subtype
Tectate; psilate. Sometimes the tectum is a tectum perforatum.
Plukenetia volubilis Linn. Plukenetia volubilis subtype; oblate.
Baker 153 PI. XII, 4 P 36 E 56 P:E 0,64.

[U] = =
ft
=
P.A.I. = 0,35.
Plukenetia abutifolia Pax et Plukenetia volubilis subtype.
(Ducke)
K. Hoffm. P = 50 E = 62 P;E =
0,80.
n n
Ducke 20619 [U] Tectum perforatum.
Eleutherostigma lehmannianum Pax et Plukenetia volubilis subtype.
K. Hoffm. P = 37 E =
47,5 P;E =
0,79.
fi n
Lehmann 5158 [K] PI. XIII, I P.A.I. =

0,35.
POLLEN MORPHOLOGY EUPHORBIACEAE 63
OF|THE
Tetracarpidium conophorum (Muell. Arg.) Plukenetia volubilis subtype.

Pax et K. Hoffm. P =
30,5 E = 44 P;E =
0,69.
n n
[= Plukenetiaconophorum Muell. Arg.] P.A.I. =

0,15 —
0,2. Margo.
Mann 1739 [U]
Versuchsanst. 9 [U]
Pterococcus corniculatus (E. Smith) Pax et Plukenetia volubilis subtype.
K. Hoffm. P =
32 E =
44 P:E =
0,66.
n n
[= Plukenetia corniculatus E. P.A.I. 0,2.

Smith] =
Koorders 41720 B [U]

Anabaenella tamnoides (Juts.) Pax et Plukenetia volubilis subtype.
K. Hoffm. P = 43 E =
49,5 P:E = 0,87.
n fi
nom. illeg. [ = Romanoa ] Trevisan P.A.I. =

0,2. Margo.
Plukenetia tamnoides (Juss.) Muell.
=
Lutz [U]
Angostyles longifolia Benth. Plukenetia volubilis subtype.
Ducke 23528 [U] P=45 Ju E = 60 P:E =
0,75.
/<
P.A.I. =
0,45. Tectum perforatum.
Astrococcus cornutus Benth. Plukenetia volubilis subtype.
Spruce 2050 [K] PI. XIII, 3 P =
36,5 n E =
50
n
P:E =
0,73.
P.A.I. = 0,4. Margo; tectum
per-
foratum.
Haematostemon guianensis Sandwith Plukenetia volubilis subtype.

Fanshawe 6016 [U] P = 29 E=40/r P:E =
0,74.
/j.
P.A.I. =
0,25. Margo; tectum
per-
foratum indistinct.
Omphalea diandra Linn. Plukenetia volubilis subtype.

Frees (Krukoff) 1893 [U] PI. XIII, 2 P =
21,5 n E =
30,5 P;E = 0.59.
n
Lanjouw and Lindeman 3164 [U] P.A.I. =

0,3.
Thunb. Plukenetia volubilis subtype;
Tragia capensis spheroidal.
Hooker Herb. [K] P = E = 17 a.
/
Plukenetia verrucosa subtype
Intectate reticulate.
(?);
Plukenetia verrucosa Smith Plukenetia verrucosa subtype; suboblate.
B.W. 993 [U] PI. XII, 1 P = 28 E = 35 P:E =

0,80.
/x //
P.A.I. =
0,3. Lumina 2-3 fi-
Apodandra buchtienii (Pax) Pax Plukenetia verrucosa subtype.

[= Plukenetia buchtienii Pax]
Krukoff 10753 [U]
Fragariopsis scandens St.-Hill. Plukenetia verrucosa subtype.
D’Alleizette 6594 [L] PI. XII, 2 P =
33,5 E = 41,5 P:E = 0,81.
n n
P.A.I. =
0,45.
Tragia stolziana Pax et K. Hoffm. Plukanetia verrucosa subtype.
Stolz 1775 [U] PI. XIII, 4 P=29/t E =
32 P:E =
0,90.
Reticulum consisting of
separate pila,
lumina P.A.I.
1-2 (i. =
0,6.
Sphaerostylis natalensis (Sond.) Croizat Plukenetia verrucosa subtype.
v.
Someren 2409 [K] P 38,5 E=43/t P:E 0,89.
=
ti =
P.A.I. =
0,6. Lumina 1-2 /i.
Sphaerostylis tulasneana Baill. Plukenetia verrucosa subtype.

D’Alleizette, Nossibe. Nov. 1906 [L] P = 35 /« E = 44 P:E =
0,84.
/I
P.A.I. =
0,3. Lumina 1-2
/i.
Tragia tristis subtype
Intectate; pilate. Pila not in a reticulum.
Tragia tristis Muell. Arg. Tragia tristis subtype.

Hassler 6645 [BM] PI. XIII, 5 P = 27 E = 40 fi P;E =
0,68.
n
P.A.I. = 0,3.
64 W. PUNT
plate xin. 1. Eleutherostigma lehmannianum; 2. Omphalea diandra; 3. Astrococcus
4. Tragia stolziana; 5. Tragia tristis; 6. hirsutum.

cornutus; Pachystilidium
Tragia geraniifolia Baill. Tragia tristis subtype.

Herter 1006 A. [U] P” 31,5 /i E
=
41,5 = P:E =
0,76.
ti
P.A.I. =
0,3. Pila ca. 1
fi.
Tragia volubilis Linn. Tragia tristis subtype.

Focke 992 [U]
Tragia ramosa Torr. Tragia tristis subtype.
Rogerson 4-6-1954 [U]
Pachystylidium type
Triporate; spheroidal.
Apertures are circular in shape and sunk down. The borders are indistinct.
Tecta psilate.
te;
The has circular of which the rims

Pachystilidium type apertures,
are indistinct. It is, however, not clear if the type belongs with
certainty to the Plukenetia configuration. The ektexine is only very
weakened and it is therefore call the

indistinctly possible to type
The is thinned. In the
inaperturate. endexine, however, clearly
author’s opinion it is better to
keep the type in the Plukenetia configura-
tion and in the Cnesmosa
not configuration.
Pachystilidium hirsutum Pax et Pachystylidium

(Blume) . ,
type.
K. Hoffmann P =
E =
29 fi. P.A.I. =
0,6-0,7.
Elbert 3340 PI. 6
[L] XIII,
Chiropetalum configuration
Tricolpate or stephanocolpate (4); shape variable.
No colpi. Colpus membrane (endexine)

Colpi long. costae persistent.
[Rarely tricolporate, but then pollen grains prolate in shape and an operculum
present.]
This configuration is distinguished from the Plukenetia

configuration
the which is
by colpus membrane, never
ruptured.
The genera of the Chiropetalum type, Ditaxis type and Argythamnia

related. Mueller and Bentham united them
type are closely (1866)
in one taxon, the Argythamnia (except Leucocroton). Pax and
genus
K. Hoffmann
(1912c, 1931) kept the genera separated, and according
to
pollen morphology this seems the right opinion.
Chiropetalum type
Tricolpate (or rarely tricolporate); spheroidal to prolate spehroidal.

Colpi wide; operculum present; costae colpi absent.
Tectate or intectate; finely reticulate.

[Rarely colpus transversalis present.]
The Chiropetalum type and the Lasiocroton

type are related by the
of and its
presence an operculum. Chiropetalum allies, however, are
finely reticulate.
From the Ekman 16425a sheet Erdtman described Leucocro-
(1952)
The Ekman sheet 12343 examined
ton
flavicans as
colpate. as
by the
66 w. PUNT
author from the Kew herbarium has These

tricolporate pollen grains.
be in
pollen grains should placed a different configuration, although
other characters (e.g. the operculum and fine reticulum) are consistent
with the Noth the transversalis

Chiropetalum type. withstanding colpus
"
the author Leucocroton

prefers to
place the pollen grains of flavicans in
the
Chiropetalum type.
Chiropetalum lanceolatum A. Jussieu Chiropetalum type; spheroidal.

Wedermann 34 PI. XIV, 2 P E 37,5 P: E 1.
[U] = =
fi ca.
P.A.I. =
0,3. Lumina ca. 1
/J,.
Intectate.
Chiropetalum canescens Phil. Chiropetalum type; prolate spheroidal.

Wedermann 773 [U] Lumina 1-2 Intectate.
fi.
Aonikena
patagonica Spego. Chiropetalum type; prolate spheroidal.
[= Chiropetalum patagonica (Spegg.) P =
34,5 n E =
30,5 P:E =
1,12.
n
O’Donn. et Lourteig] P.A.I. = 0,3. Lumina 1-2
fi.
Eyerden, Beetle and Grond 24370 [K]

PI. XIV, 4
Leucocroton flavicans Muell. Arg. Chiropetalum type; subprolate.

Ekman 13243 PI. 5 P *= E P:E
[K] XIV, 30,5 n =
25,5 =
1,19.
Colpus transversalis without costae.
P.A.I. =
0,4. Lumina ca. 1
fi.
Ditaxis
type
Tricolpate; oblate to suboblate. The pollen grains are bi-laterally symmetric.

One lobe of the three-lobed pollen grain is smaller than the other two.
Colpi long and not narrow.

very
Tectate; psilate; not reticulate.
This has Of the three lobes

type bi-symmetric pollen grains. one
is smaller than the other two. This curious shape was observed in six
of Ditaxis. candicans does these bi-

species Argythamnia not
possess
symetric pollen grains.
It could be whether other

not investigated species of Argythamnia
do show the Ditaxis
perhaps type.
Croizat (1945) did not see reason to Ditaxis and
any separate
The difference in the
Argythamnia. pollen type, however, supports
of Pax and K. Hoffmann who maintain
opinion (1931), Ditaxis
next to
Argythamnia.
Pax
Ditaxis fendleri (Muell. Arg.) et Ditaxis
type; oblate.
K. Hoffm. P =
30,5 E =
51 P:E =
0,60.
n n
Staffers 1703 [U] PI. P.A.I. 0,15.
XIV,_1 =
Ditaxis catamarcensis (Griseb.) Pax Ditaxis

type.
Brizuela 759 [U] P =
33 E = 42 P:E 0,79.
n n =
Ditaxis diversiflora Glokey Ditaxis type.

Clokey 7576 [U] Some pollen grains 4—colpate.
Ditaxis fasciculata Vahl ex A. de Juss. Ditaxis
type.
Boldingh 2733 A [U] P =
43 /i E = 50 n P:E =
0,86.
P.A.I. =
0,2 —
0,25.
Ditaxis lancifolia Schlechtend. Ditaxis
type.
Broadway 9257 [U] P.A.I. =
0,2 —
0,25.
Ditaxis salina Pax et K. Hoffm. Ditaxis type.
Pedersen 2638 [U]
plate xiv. 1. Ditaxis fendleri; 2. Chiropetalum lanceolatum; 3. Argythamnia candicans;

4. Aonikena patagonica; 5. Leucocroton flavicans; fi. Lasiocroton macrophyllus.
68 w. PUNT
Argythamnia type
Stephanocolpate; oblate spheroidal.

Colpi long, not
very narrow.
Tectate; the of the tectum reticulum is superimposed.

on
top a
Although there similarities with the Ditaxis

are some
type, shape
as well as structure differ too much for the
pollen grains to be
placed
in the same
type.
Argythamnia candicans Sw. Argythamnia type; 4 (-3)-colpate.

Boldingh 3401 B. [U] PI. XIV, 3 P =
42 E=45/t P:E =
0,93.
n
P.A.I. =
0,5.
Lasiocroton
type
Tricolpate or stephanocolpate; spheroidal to oblate spheroidal.

Colpi wide; operculum present; no costae present.
reticulum
Tectate; psilate; no present.
The difference between the

Chiropetalum type and Argythamnia type
is found in the absence of a reticulum.
According to Erdtman (1952) the pollen grains of Lasiocroton
macrophyllus are reticulate. With the aid of a phase contrast microscope

it that the of the Harris 11868
was
possible to
conclude, pollen grains
specimen are certainly not reticulate.
‘Taxonomic discussion
Lasiocroton and
Adelia,together with Leucocroton (p. 66), formed the
series
Adeliiformes (Pax and K. Hoffmann 1914, 1931) in the subtribe
Mercurialinae. These three genera are
certainly related
by the
following
characters:
1. Disc in the male as

well as
in the female flowers. This
present
disc is annular.
usually
2. Ovarium rudiment in the male flower absent.
3. Petals absent. Number of 5.

sepals usually
4. Plants usually dioecious and only growing in the New World.
Lasiocroton macrophyllus (Swartz) Griseb. Lasiocroton oblate spheroidal.

type;
Harris 11868 [K] PI. XIV, 6 P = 18,5 n E =
19,5 fi P:E =
0,95.
P.A.I. =
0,3.
Adelia ricinella Linn. Lasiocroton spheroidal.
type;
Rutten-Pekelharing [U] (3-) 4-coIpate. P = E = 33
[i.
Fuertes 833 [U] Operculum narrow. P.A.I. =
0,4-0,5.
Cephalomappa configuration
Tricolpate.
Colpi very short; costae colpi present.
Cephalomappa type
Tricolpate; oblate spheroidal.

Colpi very short; costae colpi present.
intra-reticulum
Tectate; coarse.
differs from
The Cephalomappa type completely all other types in
the Crotonoideae. Besides the very coarse reticulum the colpi are short
and bordered distinct

by costae colpi.
The genus is related those in the

Cephalomappa certainly to
Cladogynos
type (p. 93).
Cephalomappa malloticarpa J. J. Smith Cephalomappa type.

..
Herb. Lugd. Batav. 924.536. 117 [L] P 30,5 fi E

= = 32,5 p
P:E =
0,94.
PI. XV, 5 Colpi length ca. 4 /1 Lumina 2-3
.
fi.
SUMBAVIA CONFIGURATION
Tricolporate -
stephanocolporate; usually Polar axis shorter than Equatorial
axis.
Colpi wide or narrow.
Tectate or intectate; reticulate. Reticulum coarse.
The character of this is the

most
important configuration coarse
reticulum. Some in the also

species Cladogynos configuration (p. 93)
have reticulum but in that Polar axis
a
fairly coarse case
they have a
than the axis. In the Sumbavia the

larger Equatorial configuration
have Polar axis shorter than the
pollen grains usually a Equatorial
axis.
Most in this the

genera configuration belong to Chrozophoreae
Pax and K. Hoffmann These authors
(Bentham, 1880; 1912c, 1931).
split up
this tribe in groups. Bentham’s third
group
is comparable
with the subtribe of Pax and K. in which
Regulares Hoffmann,
Sumbavia, and found.
Chrozophora Caperonia are
Sumbavia
type
oblate spheroidal to oblate.

Tricolporate;
Colpus transversalis large; broad and
or long narrow.
Colpi long; costae colpi present or absent.
Intectate; reticulate. Reticulum coarse.
The of the Sumbavia and

pollen grains type are always tricolporate
have distinct reticulum.
a
They differ from the
Chrozophora type
and
and from the

Caperonia subtype by the number of colpi Philyra subtype
the reticulum and colpus transversalis.
by coarse
elongated
In 1952 Erdtman described of Monotaxis
pollen grains grandiflora
which and have membrana This
are
subprolate a
granulata. discrip-
tion is in accordance with the data found The
not
by the author.
pollen grains examined were suboblate and had no membrana
The exine is the

granulata. structure, however, quite same.
In the of Pax and K. Hoffmann and

system (1931) Thyrsanthera
Sumbaviopsis are placed next to Sumbavia. This is in with the
agreement
close of the
relationship pollen grains.
70 W. PUNT
Monotaxis differs from the other genera in the Sumbavia the

type _
by
absence of stellate hairs and the presence of a disc and an ovarium
rudiment in the male flowers. The flowers have, however, also five
in genera of the Sumbavia

petals as most
„ . . type-
.
in resembles Sumbavia and

many respects Melanolepis Sumbaviopsis.
As with these there is no
disc or ovarium rudiment in the
genera
male flower. On the other hand the female flower possesses a disc.
The have stellate hairs, monoecious and show

plants they are
similarity in the form of the leaves. Melanolepis, however, has no petals.

Baillon (1858) therefore calls Melanolepis a Sumbavia without petals.
Mueller Bentham and Pax and K. Hoffmann
(1866), (1880) see
Mallotus relation of An
as nearest
Melanolepis. comparison accurate
of the corresponding characters will have to decide whether Melanolepis

is closer related Mallotus than Sumbavia and
to to
Sumbaviopsis.
Sumbavia rottleroides Baill. Sumbavia
type; oblate spheroidal.
Merrill Sp. Blancoanae 993 [L] P =
43ju E = 46 P:E =
0,93.
PI. 2 0,6. P.A.I. 0,35.
XV, m:e = =
Costae colpi absent. Lumina 3-4

/x.
Sumbaviopsis albicans J. J. Smith Sumbavia type; suboblate.
Backer 19016 [L] P =

32 n E =
38,5 fi P:E =
0,80.
P.A.I. =
0,3. Lumina 1-2 /x.
Costae colpi.
Thyrsanthera suborbicularis Pierre Sumbavia suboblate.
ex
type;
Gagnepain P =
43,5 E =
52 P:E =
0,84.
i* n
Pierre 512 [K] PI. XV, 3 P.A.I. =
0,3. Lumina 1-2 /x.
Costae colpi absent.
Melanolepis multiglandulosa (Reinw. Sumbavia suboblate.

ex
type;
Blume) Reich.f. et Zollinger P =
32 £ =
38,5/1 P:E =
0,82.
/*
Hort. Bogor. 2127 [U] PI. XV, 1 P.A.I. =
0,3 —
0,35. Lumina 1-3

/x.
Monotaxis grandiflora Endl. Sumbavia oblate.

type;
Pritzel XI, 1900 [U] PI. XV, 4 P =
30,5 E = 36 P:E =
0,84.
/j.
m:e < 0,5- P.A.I. =
0,15.
Colpus transversalis long and narrow.
Lumina 1-2 fi.
Chrozophora type
Stephanocolporate; suboblate -
oblate.
Colpus transversalis broad, isodiametric; costae.
Colpi short; costae colpi.

Intectate reticulate. Reticulum
(?); coarse.
The lumina of the reticulum much in the

are larger Chrozophora
of the Sumbavia Besides
type than in any other type configuration.
this character the colpi are short and largethe number of colpi is
(at least six). In the subfamily of the Crotonoideae pollen grains with
than three four rather the related
more or
colpi are rare.
Only
Caperonia type has also six colpi.
Chrozophora plicata (Vahl) A. de Juss. Chrozophora type; (7-) 8 (-9)-colporate;

Feinbrun 49169 A. PI. 6 suboblate oblate.
etc. [U] XV, —
P = 60 n E = 80
ft
P:E = 0,75.
m:e = ca. 1. P.A.I. > 0,5. Lumina
4-6
fx.
Chozophora tinctoria (Linn.) A. de Juss. Chrozophora type. 6-colporate.

Faure 31126 A. [U]
plate xv. 1. Melanolepis multiglandulosa; 2. Sumbavia rotlleroides; 3. Thyrsanthera

suborbiculatus; 4. Monotaxis grandiflora; 5. Cephalomappa malloticarpa; 6. Chrozophora
plicata.
72 W. PUNT
Caperonia type
Stephanocolporate or tricolporate; oblate spheroidal -

prolate spheroidal.
Colpus transversalis broad, circular -
broad elliptic.
Colpi long and narrow; costae colpi absent.
Intectate; reticulate. Lumina fairly small. Columellae elongated.
This resembles the Chrozophora in its number of colpi

type type
The the lumina of the
(usually six). colpi are, however, longer, and
reticulum much smaller. The Philyra subtype differs from the Sumbavia
by its large circular colpus transversalis.

type
Mueller (1866) and Bentham (1880) included

Philyra in the
genus
Pax and K. Hoffmann
Argythamnia. (1912c, 1931) separated Philyra
from this genus and referred it close to
Caperonia. Pollen-morpholog-
the genus is related than and
ically more to Caperonia to Argythamnia
its allies.
Caperonia subtype
Pollen grains stephanocolporate.
Caperonia palustris (Linn.) St. Hill. Caperonia subtype; oblate spheroidal.

Lanjouw 1044 [U] PI. XVI, 4 P = 32 E = 33 P:E =
0,94.
n n
m:e ca. 1. P.A.I. =

0,3. Lumina
ca. 1
fi.
Caperonia serrata Presl Caperonia subtype; spheroidal.

de Wit 429 [WAG] P = E = 39 n. P.A.I. =
0,5.
Caperonia corchoroides Muell. Arc. Caperonia subtype.
Rombouts 358 [U]
Philyra subtype
Pollen grains tricolporate.
Philyra brasiliensis Klotzsch Philyra subtype; prolate spheroidal.

Hassler 12323 [K] PI. XVI, 2 P =
38,5 E =
35 P;E = 1,10.
fi n
m: e =
0,7. P.A.I. =
0,4. Lumina
1-2
ft.
Caryodendron type
Tricolporate; oblate spheroidal -

suboblate.
In polar view pollen grains convex triangular.

Colpus transversalis broad; costae
no
present.
Colpi narrow; no costae colpi.
Tectate, tectum perforatum; psilate. Intra-reticulum coarse, but lumina small.
The is related the Sumbavia type. The

Caryodendron type closely to
pollen grains are, however, tectate and the lumina of the intra-
reticulum are fairly small. The tectum, moreover,

is perforate. The
holes in the tectum are very

small and
only visible with
high magni-
fication.
is related the in the Sumbavia

Caryodendron not
closely to
genera
It lacks stellate indument and Bentham

type. a petals. (1880) as
well as Pax and K. Hoffmann (1914, 1931) thought of a relationship

POLLEN MORPHOLOGY OF THE EUPHORBTACEAE 73
with Alchornea. however, annular disc in the

Caryodendron possesses, an
female flower, a character never to be found in the genera of the
Alchornea but in the of the Sumbavia type.

type, always present genera
This character is also in of the Bernardia and
present genera type
On the other hand the authors cited above
Speranskia type. stated,
that in their Adenophaedra is related Caryodendron.
opinion closely to
Adenophaedra belongs pollen-morphologically to the Bernardia

type
(p. 75). The pollen grains of Caryodendron show indeed some resem-
blance with this type, but the differences are too for a com-
many
bination of and in the
Adenophaedra Caryodendron same
type.
Caryodendron orinocense Karst. suboblate.

Caryodendron type;
Krukoff 5553 [U] PI. XVI, 3 P =
29 E =
34,5 P:E =
0,84.
fi n
m:e =
0,4. P.A.I. =
0,3.
Lumina ca. 1 muri thick.
/i;
Caryodendron grandifolius Muell. Arc. Caryodendron type; oblate spheroidal.

Kuhlmann et Ducke 16598 [U] P =
31,5 n E =
33 p P:E =
0,95.
P.A.I. =
0,2.
Nealchornea type
Tricolporate; oblate spheroidal. In polar view the pollen grains circular.
Colpus transversalis broad; ora large.

Colpi short and wide.
Intectate; reticulate. Reticulum coarse.
The Polar axis shorter than the

pollen grains have a Equatorial
axis and there is also a reticulum with large lumina. For these reasons
the type is in the Sumbavia differs

placed configuration. It from the
other the three short

types by colpi.
Nealchornea is not related to discussed in other of

any genus types
the Sumbavia configuration. Its nearest allies are perhaps Hamilcoa,
and
Plagiostyles Pimeleodendron
(p. 103). Striking corresponding
characters with these genera are
the
very
short to sessile anthers and
thickened pedicels.
Pax and K. Hoffmann the in the subtribe
(1931) put genus
with Moultonianthus and

Cluytiinae e.g. Trigonostemon, Trigonopleura.
The latter three all in different and
genera are placed pollen types
in different
even
configurations.
Nealchornea yapurensis Huber Nealchornea type.

Krukoff 4967 [U] PI. XVI, 1 P =
27 E =
30 /* P:E =
0,90.
fi
m:e = ca. 1. Lumina 3-4
fi.
Bernardia configuration
Tricolporate or stephanolcoporate (except Erismanthus type). Usually Polar
axis larger than Equatorial axis.
Colpi narrow;
costae colpi.
Tecta te or in tecta
te; reticulate. Reticulum fine; lumina <2 // ( Cyttaranthus
without reticulum).
The of the Bernardia configuration are finely reticulate and

types
have Polar axis
usually a larger than the Equatorial axis. Cyttaranthus
74 W. PUNT
1. Nealchornea yapurensis; 2. Philyra brasiliensis; 3.

plate xvi.
Caryodendron orinocensis;
4. Caperoniapalustris; 5. Pseudagrostistachys africana; 6. Heterocalyx laoticus; 7. Cyttaranthus
congolensis; 8. Bernardia pulchella.
in other
only is not reticulate, but respects the pollen grains of this
resemble the in this
genus closely Speranskia type. Although placed
configuration by their fine reticulum and prolate shape, the Moul-
tonianthus
type and the Clutia
type
differ largely from the Bernardia
and the
type Speranskia type.
Bernardia type
Tricolporate; spheroidal prolate -

subprolate. Pollen grains in polar view
distinctly three-lobed.
Colpi costae colpi. P.A.I. small < 0,25.

narrow;
Intectate; finely reticulate; lumina ca. 1

fi.
The Bernardia differs from the

type Speranskia type by its distinctly
three-lobed pollen grains.
Mueller originally described Adenophaedra as a section of Bernardia.

In later
publications he raised it to a
separate genus. Bentham (1880)
and Pax and K. Hoffmann maintained the and
(1931) two
genera
even
placed them in a
far-away group to which Alchornea belongs.
Pollen-morphological study of the two
genera
showed a close relation,
it better maintain in the
so seems to
Adenophaedra vicinity of Bernardia.
Both are restricted to the New World.
genera
Bernardia pulchella (Baillon) Muell. Bernardia prolate spheroidal.

type;
P = 25,5 E =
24/i P:E =
1,07.
ft
Tessman 6078 [U] PI. XVI, 8 m:e = ca. 0,5. P.A.I. =

0,15.
Bernardia lorentzii Muell. Arc. Bernardia
type.
Schulz 1554 [U] P = 29 E =
25 P:E =
1,17.
n n
P.A.I. =
0,2.
Bernardia corensis Muell. Arg. Bernardia
(Jacq..) type.
Boldingh 665 B. [U] P = 31 E =
25 P;E =
1,24.
iu /x
P.A.I. =
0,2.
Adenophaedra minor Ducke Bernardia
type.
Herb. Rio 10386 [U] P =
28 E 27 = P:E =
1,04.
n /i
Colpus transversalis
narrow;
m:e < 0,5. P.A.I. =
0,1-0,15.
Speranskia type
Tricolporate; spheroidal —
prolate spheroidal. Pollen grains in polar view
convex triangular.
Colpus transversalis large; costae or absent.
present
Colpi narrow; costae colpi. Sometimes costae indistinct.
Tectate; intra-reticulum rarely absent

present or
( Cyttaranthus).
This looks somewhat like the Bernardia type but differs by its
type
and
polar view the shape of the colpus transversalis.
Cyttaranthus
belongs undoubtedly to this but lacks the fine reticulum.
type
Genera with found in the Old World.

a
Speranskia type are
only
They have several characters in common.
1. In the female flowers always a disc present.
2. Anthers usually pendulous; connectives frequently appendiculated.

76 w. PUNT
3. In the male flowers ovarium rudiment is

an
wanting except in
some Agrostistachys species.
4. The Asiatic The
genera have petals and an extrastaminal disc.
African lacks while
representatives frequently petals, glands are
always present on the receptaculum.

5. Stellate hairs never present.
Speranskia pekinensis Pax et K. Hoffmann Speranskia type; subprolate.

Bullock 10 Mai. 1888 [K] PI. XVII, 8 P =
28,5 E=24/t P:E = 1,16.
n
m:e =
0,5-1. P.A.I. =
0,10.
Lumina < 1
/s.
Pseudagrostistachys africana (Muell. Arg.) Speranskia type; subprolate.

33 E =
26,5 P;E =
1,24.
/i /*
Moller and Quintus 16 [U] PI. XVI, 5 m:e < 0,5. P.A.I. =
0,3.
Lumina ca. 1
fi.
Cyttaranthus congolensis Leonard Speranskia type; prolate spheroidal.

Gollens 3541 [K] PI. XVI, 7 P = 30 E =
28,5 P:E =
1,05.
n /i
P.A.I. =
0,35. Not reticulate!
Agrostistachys indica Dalz. Speranskia type; oblate spheroidal.

Stocks [U] P =
31,5 E =
33,5 P:E =
0,94.
fi n
m:e < 0,5. P.A.I. =
0,25. Lumina
1-2 of the
ft. Margo (thinning ektexine).
Agrostistachys sessilifolia (Kurz) Pax et Speranskia type; subprolate.
K. Hoffmann P =
26,5 E =
23,5 P:E = 1,12.
/j. fi
East Ind. Comp. 4739 [U] P.A.I. =

0,2-0,25.
Costae transversales. Margo (thinning
of the ektexine).
Heterocalyx laoticus Gagnep. Speranskia spheroidal.
type;
Kerr 20895 PI. 6 P E 30 P.A.I.
[K] XVI, =
=
/i.
=
0,25.
Lumina 1-2 Margo (thinning of the
/(.
ektexine).
Argomuellera macrophylla Pax Speranskia type; spheroidal.
de Wit 32 [WAG] PI. XVII, 3 P =
25,5 E =
25 P;E =
1,02.
n /i
m:e =
0,7-0,8. P.A.I. =
0,4.
Lumina ca. 1 Costae transversales
/i.
absent.
Swynnerton 119 A [BM] P = 30 fi E =

28,5 n P:E = 1,08.
In other characters as above.
Discoglypremna caloneura Prain Speranskia prolate spheroidal.

(Pax) type;
Zenker 2643 [L] P=27 E =
25 P:E =
1,09.
ii ,m
P.A.I. =
0,25-0,3. Lumina ca. 1
fi.
Moultonianthus
type
Tricolporate; spheroidal. In polar view the pollen grains circular.
Colpus transversalis narrow and small; costae.
Colpi narrow and long; costae colpi.

Intectate; reticulum fine.
The but
pollen grains of the Moultonianthus type are
finely distinctly
reticulate and therefore in the Bernardia
are placed configuration.
The view and transversalis of this differs
shape, polar colpus type
from the other in this
greatly types configuration.
Steenis Moultonianthus is related Eris-

According to v.
(1948) to
manthus in several characters. Unfortunately pollen grains of Eris-
manthus indochinensis do not

agree with those of Moultonianthus leem-
of Erismanthus will show rela-

bruggianus. Perhaps other species more
in their
tionship pollen-morphology.
Moultonianthus leembruggianus (Boerl. et Moultonianthus type.
Koord.) v. Steenis P= E =
23,5 fi. m;e < 0,5.
Sarawak 464 PI. 1 P.A.I. Lumina 1-2
[L] XVII, =
0,4. n-
Erismanthus type
Tricolpate; oblate spheroidal. In polar view pollen grains circular.
Colpi narrow.
Rim of the colpi irregularly ruptured.
indochinensis
Pollen grains of Erismanthus distinctly differ from the
There
Moultonianthus-type. are no composite apertures and the grains
Erdtman examined Erismanthus and
are tectate. (1952) oblique on
the did find in this

contrary composite apertures specimen.
Erismanthus indochinensis Gagnep. Erismanthus

type.
D’Alleizette 942. 20-4-1908 [L] P =
24 E =
25,5 P:E =
0,93.
n n
PI. XVII, 4 P.A.I. = 0,4.
Clutia type
Tricolporate; subprolate. In polar view the pollen grains triangular.

Colpi narrow; costae colpi. Margo present (thinning of the ektexine).

Tectate; intra-reticulum indistinct.
The Clutia type strongly recalls the Phyllanthus pentaphyllus su btype

in the subfamily of the Phyllanthoideae. The type differs, however,
the of the
by shape colpus transversalis. The Clutia type differs from
the Bernardia its polar view. The Bernardia
type by type is three-lobed,
the Clutia type triangular.
Taxonomical discussion
This African genus stands alone in the Euphorbiaceae. The in-
florescence and habit those of but the essential

are Phyllanthus;
characters those of the
are Chrozophorae (Bentham, 1880).
Clutia paxii Knauf Clutia type.
Crookewit 198 [WAG] PI. XVII, 2 P =
35 /i E =
27 n P:E =
1,30.
m:e < 0,5.
Clutia natalensis Bernh. Clutia type.
Wylie Jan. 1935 [U] P = 43 E = 33 P:E = 1,30.
n n
Reticulum indistinct.
Clutia abyssinica Jaub. et Spach Clutia

type.
Stolz 1876 [U] P = 52 E == 40 P:E =
1,30.
fi n
Reticulum indistinct.
Clutia rubricaulis Eckl. Sond. Clutia

ex
type.
[= C. alaternoides Linn.] P —
54 E = 41 P:E =
1,30.
fi fi
Lanjouw 165 [U] Reticulum indistinct.
Mallotus configuration
Tricolporate -
stephanocolporate (4). Polar axis

usually shorter than Equator-
ial axis.
Colpi narrow.
Tectate perforate in Blumeodendron usually not reticulate;

(tectum type);
columellae short, capita indistinct.
78 W. PUNT
1. Moultonianthus leembruggianus; 2. Clutia paxii; 3.

plate xvu.
Argomuellera macro-
phylla; 4. Erismanthus indochinensis; 5. Mallotus albus; 6. Cleidion javanicum; 7. Macaranga
spinosa; 8. Speranskia pekinensis; 9. Trewia

nudiflora.
The characters of the Mallotus

most important configuration are
the short columellae and the indistinct

capita. Besides these important
characters the reticulate.
pollen grains are usually not
- -
- - -
Mallotus
type, Blumeodendron type, Acalypha type, Alchornea type,
Discocleidion and
type Mareya type are
closely related and differ only
in minor
points.
The Ricinus
type and Pera type are connected
by the very narrow
colpi and
nearly visible columellae. They certainly belong to the
Mallotus configuration although the shape of the grains is

pollen
often prolate.
The Seidelia and Leidesia related each
type type are to
other, es-
in their exine and

pecially structure polar view.
The Microdesmis in the
type and Cheilosa type are
placed Mallotus
configuration because of their shape and short columellae. The
differences with the other types in this however,

configuration, are
considerable.
Mallotus type
Tricolporate -
stephanocolporate (4); oblate spheroidal -
suboblate. Some-
times spheroidal.
Colpus transversalis usually small; costae.
Colpi narrow;
costae colpi.
Tectate; psilate or scabrate. Rarely intra-reticulate (Adriana).
The from the other

pollen grains are
easily distinguished types.
The colpi, short (P.A.I. is least and the
are at
0,4) shape is usually
oblate spheroidal. The columellae are so short, that only with difficulty
the be from the endexine. Costae
tectum can
distinguished colpi
distinct.
In the system of Mueller genera of the Mallotus type

(1866) are
all found in his subtribe Euacalypheae. Bentham (1880) placed them
in the subtribe Pax

Acalypheae and and K. Hoffmann (1914, 1931)
in the subtribe Mercurialinae of the tribe In all three systems
Acalypheae.
the
genera are mixed up with those, closely allied to them in the
Acalypha Alchornea Blumeodendron Discocleidion

type, type, type, type
and Mareya type. Corresponding characters are:
1. A disc is wanting in the male as well as in the female flowers.
2. An ovarium rudiment is wanting.

3. inserted
Usually numerous stamens are on a convex receptaculum
(Macaranga with few has Most times
stamens no
receptaculum).
the receptaculum is glabrous. Sometimes it is provided with
glands (some species of Mallotus and Pycnocoma) or hairs ( Wetria).

4. The majority of the genera grows
in the Old World; only Avel-
lanita is from Chile.
Mallotus subtype
Pollen grains psilate.
80 w. PUNT
Mallotus albus (Roxb.) Mueix. Arg. Mallotus subtype; oblate spheroidal.

Thomson [U] PI. XVII, 5 P =
30 E =
30,5 P:E =
0,98.
n n
m:e < 0,5. P.A.I. =
0,4.
Mallotus claoxyloides (F. v. Mueller) Mallotus subtype.
Muell. Arg. P = 30 E =
32,5 P:E =
0,95.
n /u
Buysman 1874 [U] P.A.I. =

0,4.
Mallotus miquelianus (Scheff.) Pax et Mallotus subtype.
K. Hoffm. P 20 E 21 P:E
=
// =
/u =
0,95.
Rutten-Kooistra 27 [U] P.A.I. =
0,4.
Mallotus repandus (Willd.) Muell. Arg. Mallotus subtype. Too
young.
Griffith 4760 [U]
Coelodiscus lanceolatus Gagnep. Mallotus subtype.
™
D’Alleizette Jan. 1909 [L] P =

25,5 /I E =
27 /i P:E = 0,94.
P.A.I. =
0,4.
Trewia nudiflora Ltnn. Mallotus subtype.
Hohenacker 453 [U] PI. XVII, 9 P =
27/r E =
30,5 n P:E =
0,89.
Colpi short. P.A.I. > 0,75.
very
Avellanita bustillosii Philippi Mallotus subtype.
Philippi [BM] P =
25,5 E = 26 P:E =
0,98.
n n
Costae colpi indistinct. P.A.I. =

0,45.
Cordemoya integrifolia (Willd.) Baillon Mallotus subtupe.
Gover. Pires to Kew Nov. 1908 [K] P 20 E 21,5 P:E
=
n
= =
0,93.
P.A.I. =
0,7.
Coccoceras borneënse J. J. Smith Mallotus subtype; 3-4 colporate.
F. H. Endert 2078 [L] P =
37,5 E = 39 P: E =
0,96.
n
Colpi short. P.A.I. =

0,7.
very
Deuteromallotus acuminatus (Baillon) Mallotus subtype.
20 iu E =
22,5 P:E =
0,88.
fi
D’Alleizette Mad. Nov. 1908 [L] Colpi short.
very P.A.I. > 0,7.
Wetria macrophylla (Blume) J. J. Smith Mallotus subtype; 4- colporate.
(3)
Cult. Hort. Bog. IXA. 124 [U] P E 40 P.A.I.
= =
n
=
0,5.
Adriana quadripartita (Labile.) Gaudich. Mallotus subtype.
F. Mueller, Freemantle 1861 [U] P 33,5 £ 35// P;E
v. =
ft
= =
0,96.
P.A.I. > 0,5. Intra-reticulate.
Cleidion javanicum Blume Mallotus subtype.

Cult. Hort. Bog. IX A. 38 [U] P =
25,5 E —27 P:E =

0,95.
n n
PI. XVII, 6 P.A.I. > 0,7.
Cleidion angustifolium Pax et K. Hoffm. Mallotus subtype.
Le Rat 468 P 19 E 18 P:E
[P] =
fi =
/< =
0,95.
P.A.I. > 0,7.
Macaranga spinosa Muell. Arg. Mallotus subtype.
Steiner 1292 [U] PI. XVII, 7 P = E =

14,5 P:E = ca. 1.
fi
P.A.I. > 0,5.
Macaranga harveyana (Muell. Arg.) Mallotus type.
Muell. Arg. P =
17 E = 17 P:E ca.
1.
n n =
Yuncker 15980 [U]

Macaranga densiflora Wars. Mallotus subtype; P = E =
17 p.
Gjellerup 236 [U]
Pycnocoma subtype
Pollen grains scabrate.
This differs from the Mallotus

subtype subtype by the scabrae on
the In the colpi longer than those in

tectum.
Pycnocoma cornuta are
other of the Mallotus The of

pollen grains type. colpi Pycnocoma
short the others.
macrophylla, however, are as as
Pycnocoma cornuta Muell. Arg. Pycnocoma subtype.

Leeuwenberg 3003 [U] PI. XVI11, 1 P = 33,5 p
E =
35,5 p
P;E =
0,94.
m:e =
0,4. P.A.I. =
0,3.
Pycnocoma macrophylla Bentham Pycnocoma subtype.

Zenker 1251 [L] P =
30,5 E = 32 fi P:E =
0,95.
fi
P.A.I. > 0,5.
Blumeodendron type
Tricolporate; oblate spheroidal.

Colpus transversalis small and costae.
narrow;
Colpi short and costae colpi.

very narrow;
Tectum perforatum that be regarded as a coarse reticulum.
may
This type is related with the Mallotus The

closely type. tectum,
however, is The holes rather wide and it is

perforated. are
perhaps
better of reticulum. The muri of the reticulum
to speak a coarse are
thick and consist of many small columellae.
The affinities of Blumeodendron with Botryophora are

extensively
explained by Airy-Shaw (I960).
Without doubt Ptychopyxis has several characters in common
with
Blumeodendron and
Botryophora.
1. In the male flowers a disc and an ovarium rudiment are absent.
Many stamens inserted on a

receptaculum.
2. In the female flowers disc is present. 1960, states,
a (Airy-Shaw
that in
Ptychopyxis at least some
species certainly have a disc.)
Blumeodendron tokbrai (Blume) Kurz Blumeodendron type.

Achmad 216 [L] PI. XVIII, 6 P =
21/t E =
23 P:E =
0,91.
Ptychopyxis costata Miquel Blumeodendron type.
Kings collector 6740 [L] P =

18,5 E =
21 P:E =
0,88.
n n
Ptychopyxis kingii Ridley Blumeodendron type.

de Haviland et Hose 3673K [K] P = 19,5 E =
21 P:E =
0,83.
/j. /i
Botryophora geniculata (Miq,.) Beumee ex Blumeodendron type.
Airy-Shaw P =
22 E =
23,5 P:E =
0,93.
n n
Docters v. Leeuwen 7707 [L] P.A.I. = 0,5-0,6. Colpi somewhat
longer.
Acalypha type
Tricolporate stephanocolporate; oblate to suboblate. The pollen

or
spheroidal
grains quasi porate. With high magnification the undoubtedly
are
apertures are
composite apertures.
transversalis and colpus of the size; psilate.
Colpus same costae. Tectate;
The Acalypha is undoubtedly related to the Mallotus

type type.
.
... .
The dimensions of the

colpi and the colpi transversales, however, are
so small that the composite resemble pori.

apertures
Acalypha indica Linn.

Acalypha type-
PI. 4 P 14,5 E 16[ji P:E 0,90.
Dewan, 12-9-1957 [U] XVIII, =
/t
= =
Acalypha aronioides Pax et K. Hoffm. Acalypha type; 4—5 (col)-porate.

Ellenberg
1064 [U] P"= 14 pc E 16,5 fi P:E = =
0,85.
Acalypha racemosa Wallich ex Baillon Acalypha type; 3 (col)-porate.
Versuchsanst. 410 [U] P'= 12 E = 13,5 P:E =
0,89.
n /x
Acalypha platyphylla Muell. Arg. Acalypha type; 3 (col)-porate.

Killip et Garcia 33764 [U] P =
13,5 /i E =
14,5 fi P:E =
0,93.
scandens Bentham Acalypha type; 4—5 (col) porate.
Acalypha
Y. Mexia 6386 [U] PI. XVIII, 5 P'= 17,5 E =
19,5 n P:E =
0,90.
Acalypha diversifolia Jacq. Acalypha type. Too
young.
Versteeg 466 [U]

82 W. PUNT
Alchornea type
Tricolporate; spheroidal to suboblate.
Colpus transversalis with costae.
Colpi narrow; costae colpi. Operculum present.

Tectate; psilate.
In and the Alchornea is related the Mallotus

shape structure
type to
type. The pollen grains differ from the latter type by the presence
of an operculum.
Genera of this have much resemblance with those of the

type
Mallotus In the of Mueller Bentham
type. systems (1866), (1880)
and Pax and K. Hoffmann
(1931) they are mixed up with them.
Indeed the characters of this group correspond well with those
described in the Mallotus type (p. 79). There are
only a few indistinct
differences.
1. Number of is often less than 20.

stamens Neotrewia, Gavarretia and
have 60 (In the Mallotus type up to
Polyandra up to stamens
several hundreds).
in
2. Receptaculum usually flat and
peltate (not species with many
As in the Mallotus absent

stamens). type glands are or scarce
and Neotrewia).
(Alchornea
and which
Coelebogyne Lepidoturus are genera, nowadays are con-
sidered as belonging to Alchornea.
Croizat Croizat
Originally placed Adenophaedra prealtum (Croizat)
in the genus Cleidion This has its widest in
(1943 c). genus spread
Asia and was said to have some representatives in America. After-
wards Croizat’s opinion was that all American Cleidion species belong
to
Adenophaedra (1946 a). Pollen-morphological investigation shows,
that pollen grains of the species Adenophaedra prealtum on no account
resemble those of minor Ducke. The A de-

Adenophaedra diagnosis of
prealtum however, fits into the
no
phaedra , completely new genus
Polyandra described by Leal (1951). Leal moreover published a
photograph of the pollen grains of this genus, from which it is clear

that the the Alchornea like those of
pollen grains belong to
type
Adenophaedra prealtum.
Alchornea villosa (Benth.) Muell. Arg. Alchornea type; oblate spheroidal.

Bosch Proefst. F. 1152 [U] PI. XVIII, 2 P = 25 E =
26,5 P:E =
0,94.
m:e =
0,6. P.A.I. =
0,4.
Alchornea (Lour.) Muell. Arg. Alchornea
rugosa type.
Elmer 14353 [U] P=24 E=26,5 P:E 0,92.
Ju n
==
P.A.I. > 0,5.

Alchornea schomburgkii Klotzsch Alchornea
type.
Hilleris-Lambers 1922 P 24,5 P:E 1.
=
fi _E =
24,5 ji =
Alchornea triplinervia (Spreng.) Muell. Alchornea P E 27

type. = =
/i.
Arg.
Regnell III. 1066 a [U]

Lepidoturus laxiflorus Benth. Alchornea
type.
Alchornea laxiflora (Benth.) Pax P E P:E
[= = 20,5 n
=
22 /M
=
0,93.
et K. Hoffmann] P.A.I. =
0,35-0,4.
Faulkner 531 [K]
Coelebogyne ilicifolia J. Smith [= Alchornea Alchornea type.
ilicifolia (J. Smith) Muell. Arg.] P =

20 E =
23 P:E =
0,88.
p n
F. v. Muell., Rockhampton 1873 P.A.I. = 0,4-0,45.

[K] PI. XVIII, 7
Neotrewia cumingii (Muell. Pax Alchornea

Arg.) type.
et K. Hoffm. P = 21 E = 22,5 P:E =
0,93.
/x fi
Elmer 15623 [U] P.A.I. =
0,4. Operculum consists of
only one row of pila.

Gavarretia terminalis Baill. Alchornea type.
Ducke 23511 [U] P = 27, 5 E =
30,5 P;E =
0,91.
fi
P.A.I. =
0,3.
Conceveiba simulat a Steyermark Alchornea
type.
Krukoff 6643 P 19,5 E 22,5 P:E 0,88.
[U] =
n =
n
=
P.A.I. =
0,4. Scabrate.
Conceveibastrum Pax Alchornea

martianum (Baill.) type.
et K. Hoffm. P =
21,5 E =
22,5 P:E =
0,93.
/i fi
I.B.V. 23529 [U] P.A.I. =
0,4-0,45.
Veconcibea latifolia (Benth.) Pax et Alchornea type.
K. Hoffm. P 25 E=27 P:E 0,92.
lx
=
=
/u
[= Conceveiba latifolia Bentham] P.A.I. =
0,4.
Spruce 2826 [BM]
cordatum (Juss.) Baill. Alchornea
Aparisthmium type.
Y. Mexia 6273 [U] PI. XVIII, 3 P = 24 E = 30 P:E =
0,80.
/*
P.A.I. =
0,4.
Lautembergia multispicata Baill. Alchornea
type.
D’Alleizette Madag. Nov. 1906 [L] P = 21,5 E = 25 P:E = 0,86.
/x n
P.A.I. =
0,35.
Lautembergia coriacea (Baill.) Pax Alchornea sometimes 4—colporate.
type;
D’Alleizette Madag. Oct. 1906 [L]
Bocquillonia sessiliflora Baill. Alchornea type.
Balansa 274 P 21,5 E 24,5 P:E 0,88.
[L] =
n =
n
=
P.A.I. =
0,35.
Adenophaedraprealtum (Croizat) Croizat Alchornea
type.
[= Polyandra L£al?] P =
25,5 n E = 29 n P:E =
0,88.
Krukoff 6602 [U] P.A.I. =
0,45.
Discocleidion type
Tricolporate; spheroidal to suboblate.
Colpus transversalis; costae as broad as the colpi.

Colpi broad of the P.A.I.
narrow; margo present (thickening ektexine). small,
smaller than 0,2.
Tectate; from psilate to coarsely warted. The ornamentation differs strongly.
The shows resemblance the

pollen type some to
Mareya type as
well as to the Alchornea type. It differs from both

types by its curious
ornamentation. This ornamentation varies in the individual pollen
It is certain whether other specimens of Discocleidion
grains. not
will show the variation of ornamentation.

rufescens same
The differs from the the presence of

type Mareya type by costae
and from the Alchornea type by the absence of an operculum.
Discocleidion differs from the genera in the Alchornea

type by the
presence
of a disc in the female flowers and many glands inserted on
the receptaculum of the male flowers. These characters place the genus
the of the
next to
genera Mareya type.
Mueller first described Discocleidion as a section of Cleidion. Pax
84 W. PUNT
plate xvra. 1. Pycnocoma cornuta; 2. Alchornea villosa; 3. Aparisthmium cordatum;

4. 5. Acalypha scandens; 6. Blumeodendron tokbrai; 7. Coelebogyne ilicifolia;
Acalypha indica;
9. Discocleidion rufescens; 10. Kunstlerodendron sublanceolatum.
8. Mareya brevipes;
and K. Hoffmann it better raise the section

(1931) thought to to
level which in the Bernardiiformes close Bernardia

genus they place to
and reduced
Mareya with their allies. Hurusawa (1954) at last the
genus again to a section, this time of Alchornea.
Discocleidion rufescens Pax et Discocleidion type.

(Franch.)
K. Hoffmann P = 24 E=27 n
P:E = 0,89.
D’Alleizette China Mai. 1908 [L] m:e =
0,7. P.A.I. =
0,1-0,15.
PI. 9
XVIII,
Mareya type
Tricolporate; oblate spheroidal to prolate spheroidal. In polar view pollen

grains convex triangular.
Colpus transversalis elongated; no costae.
Colpi narrow; no costae colpi present.

Tectate; psilate.
The small. The is smaller than 30

pollen grains are largest axis fx.
The is from the Mallotus the
Mareya type distinguished type by
absence of costae.
In Pax and K. Hoffmann’s (1914, 1931) of this

system genera
the Bernardiiformes. the characters the

type belong to By following
differ from those in the Mallotus and Alchornea
genera type type:
1. In the female flowers a disc is

present.
2. The male flowers have often with
a receptaculum set glands.
A disc and an ovarium rudiment are wanting. (Mareyopsis is
anomalous
by the presence
of extra-staminal disc glands and the
absence of in the
glands receptaculum.)
3. Anthers often have appendiculate connectives.
4. Plants usually grow in the Old World. (,Alchorneopsis in the
New World.)
The American genus Alchorneopsis differs from the above characters
by having an ovarium rudiment and the of hairs on the

presence
receptaculum instead of glands.
Mareya bervipes Pax Mareya type; oblate spheroidal.

Zenker 1794 [L] PI. XVIII, 8 P = 21 E =
23 P:E =
0,91.
v
m:e < 0,5. P.A.I. =

0,3.
Colpus membrane granulate.
Crotonogynopsis usambarica Pax Mareya type. Too to give reliable
young
Eggeling 3826 [K] measures.
Chondrostylis bancana Boerl. Mareya type; spheroidal,

Cult. Hort. Bogor. II. I 8. [L] p = E =
25 P.A.I. =
0,3.
ji.
Kunstlerodendron sublanceolatum Ridl. Mareya type; oblate spheroidal.

Sinclair 39882 [L] PI. XVIII, 10 P = 20,5 E = 21 P:E =
0,98.
n //
P.A.I. =
0,3.
Mareyopsis longifolia (Pax) Pax et Mareya type; oblate spheroidal.

K. Hoffmann P = 18 E = 19 P:E =
0,96.
/i n
Zenker 4228 [K] P.A I. =

0,2.
Neopalissya castaneifolia (Baill.) Pax Mareya type; prolate spheroidal.

PerviI16 387 [P] P E 21 P:E
= 22 n
=
n
=
1,05.
P.A.I. =
0,25-0,3.
86 W. PUNT
Necepsia afzelii Prain Mareya type; prolate spheroidal.

P E=21 P:E
Leeuwenberg 2211 [U] =21,5 /i ft = 1,02.
P.A.I. =
0,3.
Amyrea humberti Leandri Mareya type; prolate spheroidal.
D’Alleizette Dec. 1905 [L] P =
24 /< E=21,5yu P;E =
1,13.
P.A.I. =
0,3.
oblate
Alchorneopsisfloribunda (Benth.) Muell. Mareya type; spheroidal.
P = 18 n E = 19 P:E =
0,95.
For.’Dept. 2889 [U] P.A.I. =
0,2.
trimera oblate spheroidal.
Alchorneopsis Lanj. Mareya type;
49 P 21/t E 22 P:E 0,96.
Lanjouw 1926; [U] = =
fi =
P.A.I. =
0,25.
Ricinus
type
differs from suboblate subprolate.

Tricolporate; shape to
Colpus transversalis narrow; no costae.
Colpi very narrow; no costae colpi present.

visible in
Tectate; psilate. Columellae very short, nearly some
species.
The resemble the Pera but the Ricinus

pollen grains type, type
misses in the It differs from the latter
costae as Mareya type. type by
and visible columellae.
narrow
colpi nearly
The genera in this in

type fall apart two
groups. The first group
includes Homonoia and Lasiococca. These genera remarkable

Ricinus, are
for the number of anthers crowded branched

large on
repeatedly
filaments.
The second group comprises Koilodepas only, which genus occupies

isolated in the
an position family (Airy-Shaw, 1960). Croizat (1942b)
as well as Airy-Shaw reduced Nephrostylus Gagnepain and Calpigyne
Blume to Koilodepas. According to Croizat, Calpigyne hainanense and
In the of Airy-Shaw,
Nephrostylus poilanei are synonyms. opinion
the Pollen
Nephrostylus poilanei belongs to species Koilodepas longifolium.
grains of both
species are practically identical and show great structural
resemblance with bantamense.

Koilodepas
Ricinus communis Linn. Ricinus spheroidal.

type;
v.
Heerdt 531 [U] P = 32 E =
32 P:E ca. 1.
p p
Versteeg 243 [U] PI. XIX, 3 m: e < 0,5. P.A.I. =
0,2.
Lasiococca symphylliifolia (Kurz) Ricinus spheroidal,
type;
D. Hook. p E 23 P.A.I. 0,45.

J. = =
fi. =
Hort. Bogor. Cult. 6-4-1887 [L]

Homonoia javensis (Blume) Muell. Arg. Ricinus type; prolate spheroidal.
Hort. Bogor. IX A. 60 PI. XIX, 2 P 22,5 E 2l/x P:E 1,08.
[U]
= =
=
n
P.A.I. =
0,5. Costae transversales?
Homonoia riparia Lour. Ricinus
type; oblate spheroidal-suboblate.
Heifer 4746 [U] V =
2bn E = 29 P:E = 0,88.
Koilodepas bantamense Hassk. Ricinus
type; spheroidal-suboblate.
Hort. Bogor. IX C. 36 [U] Pi. XIX, 4 P =
20 fi E = 24 n P:E = 0,84.
m:e < 0,5. P.A.I. =
0,3.
Koilodepas hainanense (Merr.) Croizat Ricinus
type; subprolate.
[ =
Calpigyne hainanense Merr.] P = 21 /x E = 17,5 ft P:E =
1,18.
S. K. Lau 3542 PA.I. =0 3.
[P]
[? =
Nephrostylus poilanei Gagnep. P = 20,5 /<’ E =
17,5 /x
P:E =
1,17.
D’Alleizette, Annam 11-4-1907 [L] P.A.I. =

0,3.
plate xix. 1. Leidesia procumbens; 2. Homonoia javensis; 3. Ricinus communis; 4. Koilodepas

bantamense; 5. Chaetocarpus castanocarpus; 6. Pera bicolor; 7. Seidelia triandra; 8. Micro-
desmis casearifolia.
88 w. PUNT
Pera
type

Colpi narrow;
costae colpi narrow but distinct.

very
The Pera type has the visible

same very narrow
colpi and nearly
columellae as the Ricinus but there are distinct costae colpi and
type,
costae transversales
present.
Pera bicolor (Klotzsch) Mueix. Arg. Pera

type.
Ule 8409 PI. XIX, 6 25,5 E 24 P:E 1,07.
[U] P =
n
=
yu =
m;e < 0,5. P.A.I. =

0,3.
Pera glabrata (Schott) Baill. Pera
type.
Reitz 4548 P 19 E 18 P:E 1,06.
[U] =
n
=
n
=
P.A.I. =
0,5.
Seidelia type
Tricolporate; prolate spheroidal. In polar view the pollen grains distinctiy

three-lobed.
Colpi narrow; no costae colpi.

The of this three-lobed and have

pollen grains type are distinctly
costae transversales.
Seidelia triandra (E. Meyer) Pax Seidelia type.
MacGregor Mus. 1254 [K] PI. XIX, 7 P = 17,5 n E = 16,5 /s

P:E = 1,06.
m:e
=
0,6. P.A.I. = 0,25.
Leidesia
type
Tricolporate; oblate spheroidal. In view the pollen grains distinctly

polar
three-lobed.
Colpus transversalis narrow and long; no costae.
Colpi narrow; no costae colpi. Narrow operculum.

The Leidesia resembles the Seidelia type in its polar view and
type
exine It the absence of and the
structure.
differs, however, by costae
addition of a narrow operculum.
Leidesia and Seidelia are Mercurialis-like plants which Bentham (1880)

well Pax and K. Hoffmann enumerated close
as as (1914, 1931)
together with Mercurialis. This relationship, however, is not confirmed
by pollen-morphological results.
Leidesia procumbens (Linn.) Prain Leidesia

type.
Splitgerber [L] PI. XIX, 1 P =

19 E =
20,5 P:E =
0,92.
ft n
m:e < 0,5. P.A.I. = 0,3.
Microdesmis
type
Tricolporate; spheroidal to oblate speroidal. In polar view pollen grains

convex
triangular.
Colpus transversalis isodiametric; m:e =
1; ora large.
Colpi narrow; costae colpi broad and indistinct or absent.
Columellae
Tectate; psilate. short.
The Microdesmis is characterised by the isodiametric

type colpus
transversalis and the large ora.
Bentham discussed the genera of this in tribes,

(1880) type
two
Galearieae and Gelonieae. Microdesmis, Galearia and Pogonophora occurring

in the Galearieae have characters both of the Phyllanthoideae and
Crotonoideae. Pollen grains of these genera are, however, not at all
with those in the

comparable Phyllanthoideae.
Pierre’s Pax (1899), he described
description being unknown to
Pierre in Microdesmis Microdesmis

Centroplacus glaucinus as paniculata.
Pax, however, did have female flowers his Pierre
not at
disposal.
(1899) and Gilg (1908), on the other hand, studied female material
and found 2 ovules locule, for which the had be

per reason
plant to
placed in the subfamily of the Phyllanthoideae. The structure of the
male flowers, however, resembles that of Microdesmis to such an extent,

that it is by no means that Pax reckoned this species to
strange
Microdesmis. Since it is evident that the pollen grains of Centroplacus
be from those of Microdesmis, the of
cannot distinguished correctness
placing the in the subtribe Antidesminae

genus (Pax 1931) seems
Two conclusions
questionable. are possible:
1. link between the

Centroplacus forms a sub-family Phyllanthoideae and
Crotonoideae. On one side we find the characters that suggest a
close Microdesmis, while the

relationship to Phyllanthoideae-
character of the two ovula per locule in the ovary stands on the
other side.
2. Centroplacus has been described on the basis of material belonging

The is dioecious, that the of
to two species. plant so possibility
an erroneous combination of plants should not be overlooked.
A renewed examination of the known will

accurate material,
possibly remove the doubt.
Microdesmis subtype
Pollen grains small. Smaller than 20 fi.

Costae colpi absent.
Microdesmis casearifolia Planch. Microdesmis subtype; oblate spheroidal.

Thorenaar T. 50 [U] PI. XIX, 8 P =
15,5 E =
17,5 P:E =
0,89.
// ft
P.A.I. =
0,3.
Microdesmis zenkeri Pax Microdesmis subtype.
de Wit 4 [WAG] P = 18 E = 19 P:E =
0,95.
n //
P.A.I. =
0,5.
Centroplacus glaucinus Pierre Microdesmis subtype.
Tester 1922 [K] P = 14 E =
15,5 n P:E =
0,90.
Zenker 1761 [K] P.A.I. =
0,3.
Galearia dognaiensis Pierre ex Gagnep. Microdesmis subtype; suboblate.
Clemens 3528 [U] P =

14,5 E =
17,5 P:E = 0,83.
n /u
P.A.I. =
0,35.
Galearia filiformis (Blume) Pax Microdesmis subtype.
Blume, Java [U] P =
16,5 E = 19 P:E =
0,89.
ft fi
P.A.I. =
0,3.
90 w. PUNT
Chaetocarpus subtype
Pollen grains large. Larger than 30 //.

Costae colpi broad but indistinct.
Chaetocarpus castanocarpus (Roxb.) Chaetocarpus subtype; spheroidal.

Th WAITES P=E=29/< m:e ca. 1.
Elmer, Borneo 21064 [U] PI. XIX, 5 P.A.I. =

0,3.
Chaetocarpus schomburgkianus (O. Kuntze) Chaetocarpus subtype.
Pax et K. Hoffm.
For. Dept. 2519 [U]

Trigonopleura malayana J. D. Hook. Chaetocarpus subtype; oblate spheroidal.
Bosch Proefst. 83 E. IP. 1031 [L] P =
35 E = 38 P:E =
0,92.
n n
PI. 1 P.A.I. 0,2.
XX, =
Pogonophora schomburgkiana Miers Chaetocarpus subtype; oblate spheroidal.

B. W. 6015 [U] P =
42 E = 45 P:E = 0,93.
fi n
Krukoff 8645 [U] P.A.I. =

0,4—0,45. Tectum perforatum.
Cheilosa type
Tricolporate; suboblate to oblate spheroidal. In polar view pollen grains

convex triangular.
Colpus transversalis; costae.
Colpi narrow; indistinct costae colpi.

Tectate; echinate or micro-echinate. Columellae short.
The Cheilosa the characters of the

type possesses
most
important
Mallotus and is therefore in this
configuration placed configuration.
the
The type differs, however, distinctly by spines on the tectum.
' ‘ ■ ■■
fn Cheilosa these
spines are larger than 1 pt '
(echinate); 'in Neoscortechinia
smaller than 1 the nomenclature of Faegri and

/j. . According to
Iversen the of Neoscortechinia scabrate. The author

pollen grains are
would like call the small Erdtman

to spines microechinate according to
(1952). Echinate pollen grains are rarely found in the subfamily of
the Crotonoideae, in the Croton

only configuration.
discussed Cheilosa in the the

Bentham (1880) Acalypheae, although
male In
calyx is divided into slightly imbricate lobes. the system of
Pax and K. Hoffmann (1931) the genus is found in the tribe Gelonieae,
which also Neoscortechinia

to belongs. The pollen grains of both genera
this is not confirmed
are
undoubtedly related. Unfortunately by other
characters.
Cheilosa montana Blume Cheilosa suboblate.

type;
Korthals, Java [L] PI. XX, 9 P =
22,5 E =
29/* P:E =
0,78.
n
m;e = 0,5. P.A.I. = 0,5-0,6.
Echinate; echinae 1-2
ft.
Neoscortechinia arborea (Elmer) Pax et Cheilosa oblate spheroidal.
type;
K. Hoffm. P = 21y« E = 22,5 P:E =
0,93.
n
Hort. Bogor. IX A. 6a. [U] PI. XX, 8 P.A.I. =
0,3. Scabrate (micro-
echinate).
Claoxylon configuration
Tricolporate or stephanocolporate (4, rarely 5).

Colpi narrow.
Tectate; capita of the columellae distinct.
Pollen grains not reticulate.

plate xx. 1. Trigonopleura malayana; 2. Athroandra mannii; 3. Mercurialis annua;

4. Claoxylon cuneatum; 5. Cladogynos orientalis; 6. Cephalocrotonopsis socotrana; 7. Amperea
xiphoclada; 8. Neoscortechinia arborea; 9. Cheilosa montana.
92 W. PUNT
The Claoxylon configuration is related to the Mallotus configuration.

The capita of the columellae, however, are distinctly visible.
Claoxylon type
Tricolporate or stephanocolporate; oblate spheroidal-prolate spheroidal.

Colpus transversalis small; costae but sometimes indistinct.
present
Colpi costae colpi absent or indistinct.
narrow;
Tectate; psilate or scabrate. Columellae coarse, capita distinct.
This type is in some characters comparable with types of the
Mallotus configuration. The small spheroidal pollen grains resemble,

in their shape, those of the Mareya type (p. 85). As in the Mallotus
type (p. 79) more than three composite apertures can be present.
the type in the Bernardia is of
Placing configuration (p. 77) out
question
as a reticulum is always absent.
- - - -
The ' is the distinct of the

Claoxylon type distinguished by capita
columellae, which form a coarse and inordinate
pattern.
Genera this all the

belonging to type are readily recognised by
and attachment of the anthers. Other
shape corresponding characters
are:
1. A disc in the female flowers.

present
2. In the male flowers ovarium rudiment is absent.

an
always
Usually an extrastaminal disc is also absent, but glands are
inserted the In
on receptaculum. Discoclaoxylon receptaculum
in
glands fails, but this genus an extrastaminal disc is present.
3. Plants dioecious and in the Old World.

are usually grow only
Erythrococca stolziana Pax et K. Hoffm. Claoxylon spheroidal.

type;
Stolz 1556 [U] P = E =
22 m:e< 0,5.
n-
P.A.I. 0,5.
>
Athroandra mannii (J. D. Hook.) Pax et Claoxylon. oblate spheroidal-sub-

type;
K. Hoffmann oblate. 5 (—4) colporate.
[= Erythrococca mannii (J. D. Hook.) P =
30 E = 33,5 P:E =
0,89.
n n
Prain]
Mann 260 [U] PI. XX 2
;
Athroandra africana (Baill.) Pax et Claoxylon prolate spheroidal.

type;
K. Hoffmann P =
22,5 E=21 P:E =
1,07.
ft ft
[= Erythrococca africana (Baill.) Prain]

Zenker 453 [U]
Claoxylon cuneatum J. J. Smith Claoxylon type; spheroidal.
Versteeg 1752 PI. XX, 4 P E 21
[U] = —
fi.
Claoxylon tumidum J. J. Smith Claoxylon type; spheroidal.

Versteeg 1708 [U] 3— (4) colporate.
P = E = 19 P.A.I. =
0,4.
/t.
Psilate to scabrate.
Micrococca mercurialis (Linn.) Benth. Claoxylon type; prolate spheroidal.

D’Alleizette Mad. 1906 [L] 3- colporate.
(4)
P =
27 E =
25 P:E =
1,08.
n n
P.A.I. =
0,3.
Discoclaoxylon hexandrum Claoxylon type; spheroidal.

(Muell. Arg.)
Pax et K. Hoffm. P -
E = 21 P.A.I. =0,3.
n-
Zenker 35 [U]
Mercurialis
type
Tricolporate; subprolate to prolate.

Colpus transversalis small; costae as long as the costae colpi.
Colpi costae colpi. Operculum present but consisting
narrow; very narrow,
of only one row of columellae. Tectate; psilate. Columellae distinct.
There is a certain relation between the Mercurialis type and the
Claoxylon type. The two

types have distinct capita; both are not
reticulate and small colpi transversales. The Mercurialis

possess
- .
type
.. .
.
differs the of which is and in-

by presence an
operculum, narrow
and the
distinct, by distinctly prolate shape of the pollen grains.
Shape and attachement of the anthers show much similarity with

the genera of the Claoxylon group.
For that reason Bentham (1880)
placed Mercurialis in the immediate vicinity of Claoxylon, Erythrococca,
etc. Although the Mercurialis differs somewhat from the Cloxylon
type
there is for the author share the
type, enough affinity to opinion of
Bentham.
Mercurialis annua Linn. Mercurialis type; subprolate.

Fresh Z.-Limb. PI. 3 P E P:E
material, XX, =
25,5 y
=
22,5 y
=
1,16.
m:e < 0,5. P.A.I. =
0,4. Operculum
indistinct, consisting of some
scattered
pila on the endexine (membrana

granulata).
Mercurialis reverchonii Rouy Mercurialis
type.
Faure 18-6-1931 [U] P =
28,5 E =
22 P:E =
1,30.
/j. /j.
Mercurialis tomentosa Linn. Mercurialis type.

1317 [U] P 40 E 28,8 P:E 1,40.
Leeuwenberg =
n
=
/i
=
Cladogynos configuration
Tricolporate.
Colpi narrow;
costae colpi.
Tectate; columellae distinct. Exine thick.
The Cladogynos includes those pollen grains with

configuration
distinct columellae that bear always a tectum or tectum
perforatum.
These columellae be in reticulum but
can
arranged a more
frequently
do form The
they not a regular pattern. pollen grains are
usually
the axis is least 25 and often up 50
large; largest at
/i
to
(x.
Cladogynos type
Tricolporate; spheroidal prolate ( Epiprinus oblate spheroidal).

Colpi narrow;
costae colpi.
Tectate or tectum perforatum. Columellae distinct.
Exine thick. Some species intra-reticulate.
The character of this

most striking type is the distinct, sometimes
even tall, columellae. The pollen grains are

large and generally they
have a P:E larger than 1.
94 W. PUN'I
Pax and K. Hoffmann of this

By (1914, 1931) most
genera type
in the series of their sub tribe
are placed Cladogyniformes Mercurialinae,
which also Mallotus and Alchornea The be
to
belong. group can
distinguished as follows:
1. An ovarium rudiment present in the male flowers while a disc
is absent. The few inserted

stamens are not on a receptacle.
2. Male flowers usually in a
capitule.
3. Anthers pendulous.
4. Stellate hairs always present.
5. Plants are monoecious and grow in the Old World.
Cladogynos orientalis Spanoghe

Zipp ex Cladogynos type; prolate spheroidal.
Pierre 6213 [L] PI. XX, 5 P =
42/i E = 40 P:E =
1,05.
/r
Colpus transversalis narrow. Tectum
perforatum. P.A.I. =
0,3. Intra-reti-
culate; lumina 1-2

/s.
Cephalocroton cordofanus Hochst. Cladogynos type; subprolate.

D’Alleizette Nubie, Aug. 1909 [L] P = 49 E = 42 P;E =
1,17.
n p
Colpus transversalis broad; m:e >0,5.
P.A.I. = 0,3. Intra-reticulate.
Cephalocrotonopsis socotrana (Bale, f.) Pax Cladogynos type; prolate spheroidal.

Schweinfurth 594 [K] PI. XX, 6 P = 46 E =
42,5 fi P:E = 1,09.
P.A.I. =
0,25. Intra-reticulate.
Symphyllia siletiana Baill. Cladogynos type; prolate spheroidal.

Hort. Bog. Cult. [L] P =
27 E = 25 P;E =
1,07.
ii n
P.A.I. =
0,25-0,3. Not reticulate.
Adenochlaena leucocephala Baill. Cladogynos type; prolate spheroidal.

Hildebrandt 3258b [L] P =
53 E=48/< P:E =
1,10.
/u
m:e =
0,4. P.A.I. =
0,3.
Not reticulate.
Epiprinns poilanei Gagnep. Cladogynos type; oblate spheroidal.

Clemens 3058 [U] P =
32 E = 35 P:E 0,91.
ft n —
P.A.I. ==
Amperea type
Dicolporate or tricolporate. Shape differs with the number of colpi from
oblate spheroidal (dicolporate) to prolate spheroidal (tricolporate).

Colpus transversalis costae.
narrow;
Colpi narrow
and long; costae colpi. P.A.I. small to 0.
Tectate; psilate. Exine thick. Intrareticulate or not reticulate.
The Amperea-type shows much similarity with the Cladogynos t

>T )e -
T-.
the thick exine and the tall columellae The

Especially are striking.
much
P.A.I., however, is smaller. Some pollen grains are even syn-
colpate.
Amperea belongs to those Australian genera, that were kept separate

on account of their narrow cotyles (Stenolobae). Should this group as
such be abolished, then, the of Pax

according to statement (1924),
this has be in the Mercurialinae. This
placed group also
to
genus
comprises the genera

of the Cladogynos type.
Amperea xiphoclada (Sieb. et Spreno.) Amperea type.

Druce 2- P:E =
0,90.
Constable 26475 [U] PI. XX, 7 3- P:E =
1,02.
P =
27-29 E =
28-30
ft fi.
P.A.I. 0,15-0.
=
m:e < 0,5.
Not reticulate.
Amperea spartioides Brongn. Amperea type; spheroidal.

Tasmania [U] P = E =
26 Intra-reticulate.
ft.
Pseudocroton type
Tricolporate; prolate spheroidal to subprolate.

Colpus transversalis; costae?
Colpi costae col pi?

narrow;
Tectate; psilate. Not reticulate.
The
pollen grains of the sheet examined were not mature, so it is
difficult to put them in the proper There is some resemblance
type.
with the Speranskia type (p. 75). A reticulum, however, is absent.
As distinct columellae it best in

are
present, seems to
place this type
the Cladogynos configuration.
Bentham (1880) as well as Pax et K. Hoffmann (1931) discuss
Pseudocroton in the
Chrozophoreae. They accept relationship with
and allies. The grains have with
Argythamnia its
pollen some
affinity
the Speranskia Although the genera of this the
type. type belong to
Chrozophoreae, they are only found in the Old World, while Pseudocroton
in Central America. Affinities with the
grows genera of Cladogynos
configuration are few.
Pseudocroton tinctorius Muell. Arg. Pseudocroton

type.
O. Kuntze 1808 [K] Too to give reliable measures.
young
Hippomane configuration
Tricolporate; pollen grains in polar view circular or three-lobed.
Never convex triangular or triangular.

Colpus transversalis frequently short; sometimes narrow elongated.
Colpi usually long. A is nearly always
narrow; margo present.
Intectate or more often tectate. Columellae distinct, frequendy large.
It is define this with

not
possible to
configuration one or two
characters. The
exceptionless pollen grains are, however, easily
combination of several The
recognised by a typical characters. most
important ones are: three-lobed polar view; margo; distinct, sometimes

and If
large, columellae short colpi transversales. one of these striking
characters is absent the combination of the others still the
places
pollen grains immediately in this configuration.
The genera of this configuration can be divided into two

groups.
The first group comprises genera
of the tribe Hippomaneae of Pax and
K. Hoffmann 1931). In 1880, Bentham stated of his subtribe

(1912b,
96 W. PUNT
“This is of the natural subtribes of Crotoneae”.

Hippomaneae: one most
Indeed there is doubt of close between the

not
any a
relationship
genera.
Pachystroma is added to the first group. Evidently on technical

~
grounds Bentham added Pachystroma to the group of the Adrianeae with
Manihot, Adriana and Cephalocroton, three genera now

placed in three
different groups of Pax and K. Hoffmann

plants. (1919d, 1931)
separated the monotypic genus into the tribe Pachystromateae, although
they were aware of some affinity with the Hippomaneae. The well-
developed calyx of the male flower, which is never

present
in the
them
Hippomaneae, led to the decision to separate the genus. There are,
however, more characters in favour of a fusion than of a separation.
Pollen morphology is one of the data in favour of fusing.
The second group includes all the genera of the Euphorbieae with
the addition of Hura. It is remarkable that the typical Euphorbieae,

well characterised which
so
by their cyathium, possess pollen grains
can hardly separated from those of the Hippomaneae. Pax (1924) when
“
discussing the phylogenetic relations of the

Euphorbiaceae, pointed
already to this relationship. He noticed, e.g., the unbranched laticife-
vessels which both Pollen

rous taxa
possess. morphology is another
character the of Pax.
to support opinion
Hippomane type
Tricolporate; shape differs from subprolate to suboblate. Pollen grains in
polar view circular or three-lobed.
Colpus transversalis small (rarely elongated); costae.
Colpi and colpi. Usually

narrow long; costae
margo present .
P.A.I. is small (smaller 0,3) in species of Mabea.

except some
Tectate; psilate. Columellae sometimes tall.
The differs from the hirta its

Hippomane type Euphorbia type by
the is than the
narrower margo. Usually margo narrower
colpus
transversalis. In other respects both types can hardly be separated.
Pollen of the Omalanthus be
grains nutans subtype can separated
from the Hippomane subtype by the deeply three-lobed pollen grains.
Some species in the Hippomane type have an intra-reticulum. They
do belong altogether the Dichostema
not to
type (p. 100) as
the
upper
the columellae is fused in all thus
part of directions, forming a tectum.
Hippomane subtype
Pollen grains in polar view circular or
slightly three-lobed.
Hippomane mancinella Linn. Hippomane subtype; prolate spheroidal.

Buysman 1856 [U] PI. XXI, 3 P 32,5 /j. E
=
30,5 n P:E 1,06. =

=
m:e < 0,5. P.A.I. =

0,25.
Omalanthus populneus (Geisel) Pax Hippomane subtype.
Docters v. Leeuwen 1716 [U] P = 40 fi E=34/i P:E =
1,17.
P.A.I. =
0,15-0,20.
Mabea piriri Aube. Hippomane subtype.
B. W. 6296 [U] P = 50 ju E = 44 P:E = 1,14.
n
P.A.I. 0,20-0,25. Not
= reticulate.
Mabea taquari Aubl. Hippomane subtype.
Pulle 67 [U] P =
43,5 fi E = 41 P:E = 1,05.
n
Mabea caudata Pax et K. Hoffm. Hippomane subtype.

v. Emden 18-9-1931 [U] P = E = 70 P.A.I. =
0,4-0,3.
p.
Not reticulate.
Mabea fistulifera Mart. Hippomane subtype.
Y. Mexia 4473 P E 70 P.A.I.
[U] = =
/i. =
0,3.
Indistinct intra-reticulate.
Tectum perforatum.
Actinostemon lanceolatus Saldanha Hippomane subtype; oblate spheroidal.
'
Y. Mexia 4987 P 33,5 E 35 P:E 0,94.

[U] =
n
=
n
=
P.A.I. =
0,25. Margo absent. Costae
indistinct.
Gymnanthes lucida Swartz

Hippomane subtype; oblate spheroidal.
Jamaica, Hohsle [U] P =
25 E = 26 P:E = 0,96.
n n
P.A.I. =
0,25. Margo absent. Colu-
mellae short.
Glyphostylus laoticus Gagnep. Hippomane subtype.

Lakshnaka 1352 P E P:E
[K] =
36,5 n
= 34,5 /x
=
1,06.
P.A.I. =
0,25. Colpus transversalis
elongated. Columellae tall.
Spirostachys venenifera (Pax) Pax Hippomane subtype.

[= Excoecaria venenifera Pax] P = 22 E = 23 P:E =
0,96.
n n
Graham 1464 [K] PI. XXIII, 3 P.A.I. 0,25. absent.
=
Margo
Excoecaria bicolor Hassk. Hippomane subtype.
Hort. Bogor. IX C. 77 [U] PI. XXII, 4 P = 38 E = 36 P:E =
1,05.
n ft
P.A.I. = 0,25.
Stillingia Pax et K. Hoffm. Hippomane subtype.
gymnogyna
Wiggins 1351 [U] P 35,5 E 33 P:E

=
/u =
fi
=
1,08.
P.A.I. =
0,2.
Sapium longifolium (Muell. Arg.) Huber Hippomane subtype.
Pedersen 72 P 55 E 43 P:E
[U] =
n
=
n =
1,26.
P.A.I. =
elongated. Costae equatoriales or
almost equatorial.
Colliguaya integerrima Gill, et Hook. Hippomane subtype.
Donat 167 [U] P =
35,5 fi E = 34 P:E == 1,04.
n
P.A.I. = 0,2. Margo indistinct;
narrow.
Omalanthus nutans
subtype
Pollen grains in polar view distinctly three-lobed.
Omalanthus nutans (Forst.) Pax Omalanthus nutans

subtype.
Yuncker 15356 [U] PI. XXI, 8 P =
35 n E —
31,5 fi P:E =
1,11.
P.A.I. =
0,15.
Mabea indorum Sp. Moore Omalanthus nutans subtype.
Krukoff 1084 [U] P = 45 E =
36,5 P:E =
1,22.
n n
P.A.I. = 0,2-0,25.
Actinostemon concolor (Spreng.) Muell. Omalanthus nutans subtype.
P 30 E 34 P:E
=
n =
/t
=
0,88.
Bornmueller 536 [U] PI. XXII, 3 Columellae short.
Gymnanthes jamaicensis (Britten) Urban Omalanthus nutans subtype.

Harris 10936 [U] P E 22 absent.
= =
ft. Margo
Sebastiania chamaelea (Linn.) Muell. Arg. Omalanthus nutans subtype.
Hohenacker 830 [U] P 33/< P;E
= =
1,14.
P.A.I. =
0,2.
Sebastiania corniculata (Vahl) Muell. Arg. Omalanthus nutans subtype.
Frees 11761 [U] P E 40 P.A.I.
= =
n- =
0,15-0,2.
Macguire and Stahel 22758 [U]
Sebastiania scottiana Muell. Arg. Omalanthus nutans subtype.
Bertoni 1714 [U] P E 27 P.A.I.
= =
n. = 0,2.
98 W. PUNT
plate xxi. 1. Stenadenium spinescens; 2. Elaeophorbia drupifera; 3. Hippomane mancinella;

4. Dichostema glaucescens; 5. Pachystroma longifolium; 6. Senefeldera macrophylla; 7. Tetra-
plandra gibbosa; 8. Omalanthus nutans.

pollen morphology of the euphorbiaceae 99
Duvigneaudia inopinata (Prain) Leonard Omalanthus nutans subtype.

Zenker 217 P 23 E 20 P:E 1,16.
[U] =
/j.
=
/i =
P.A.I. =
0,2.
Maprounea brasiliensis St.-Hill. Omalanthus nutans subtype.
Williamson-Assis 8047 [U] P =
33,5 E =
30,5 P:E =
1,11.
/j. n
P.A.I. =
0,2.
Maprounea guyanensis Aubl. Omalanthus nutans subtype.
B. W. 5393 [U] P = 24 n E =
20 P:E = 1,20.
n
P.A.I. = 0,2.
Excoecaria philippinensis Merrill Omalanthus nutans
subtype.
Elmer, Palawan 13124 [U] P = 32/* E = 32/i. P.A.I. = 0,2-0,25.
Sapium ellipticum (Hochst.) Pax Omalanthus nutans subtype.
Croockewit 728 [WAG] P = E = 30 n. P.A.I. =
0,2.
Stolz 621 [U] P = 40 E = 34 P:E = 1,18.
n
Sapium sebiferum (Linn.) Roxb. Omalanthus nutans subtype.

Lilnee, Japan [U] P = 46 p.
Sapium aubletianum (Muell. Arg.) Huber Omalanthus nutans

subtype.
B. W. P 46
2141 [U] =
p.
Sapium montanum Lanj. Omalanthus nutans subtype.

B. W. 5889 [U] P =
52
p.
Sapium klotzschianum (Muell. Arg.) Omalanthus nutans subtype.

Huber P = 54
p.
Slabel, Oct. 1944 [U]

Grimmeodendron eglandulosum (A. Rich.) Omalanthus nutans subtype.
Urban P = 28 E = 26 P:E =
1,08.
fi n
Curtiss 190 [K] P.A.I. =
narrow elongated.
Bonania cubana A. Rich. Omalanthus nutans subtype (?).
Howard 5774 [U] P = 27 E =
25,5 P:E = 1,06.
/z p
P.A.I. =
0,2.
Bonania domingensis (Urb.) Urb. Omalanthus nutans subtype.
Fuertes 813 [U] B =
22 p E = 22 P.A.I. = 0,25.
n.
Columellae small.
Adenopeltis colliguaya Bert, ex Juss. Omalanthus nutans subtype.

Andreas 834 [U] P =
34,5 E =
33,5 P:E =
1,03.
n n
Dalembertia populifolia Baill. Omalanthus nutans subtype.

Hinton 4353 P 41 E 36 P:E 1,12.
[K] = =
n =
P.A.I. =
0,15-0,2. Intra-reticulate.
Tetraplandra gibbosa Pax et K. Hoffm. Omalanthus nutans subtype.

Glaziou 5613 [K] PI. XXI, 7 P = 30 n E =
27 P:E =
1,11.
n
P.A.I. = 0,15.
Ophthalmoblapton macrophyllum F. Allem. Omalanthus nutans subtype.
Glaziou 17752 [K] P = 42 E = 39 P:E = 1,08.
n /i
P.A.I. =
0,2.
Algernonia brasiliensis Baill. Omalanthus nutans subtype.
H. F. Hance 1887 [K] Young.
Pachystroma type
Tricolporate; spheroidal. Pollen grains slightly three-lobed.
Colpus transversalis broad elliptic; costae thick.
Colpi and short; costae colpi. No

narrow
margo present.
Tectate; psilate.
Pollen grains of this are strongly related to those of the

type
Hippomane The colpus transversalis, however, is broadly elliptic
type.
and the P.A.I. is much larger.
100 w. PUNT
Pachystroma longifolium (Nees) Pachystroma type.

I. M. P 41 E 41 P:E 1.
Johnston =
n
=
ft ca.
Krug and Urban 1367 [L] PI. XXI, 5 Length of colpi differs; mostly short,
sometimes long. Costae thick.
hirta
Euphorbia type
Tricolporate; subprolate to oblate spheroidal. Pollen grains in polar view
three-lobed.
Colpi broad. Sometimes operculum is

narrow; margo an
present.
psilate. Columellae distinct.
Tectate;
The this be
pollen grains belonging to
type can hardly separated
from the The is broader; broad
Hippomane type. margo mostly as
the If in the future

as colpus transversalis. The colpi are narrow. more
examined it will
Euphorbiaceae species are pollen-morphologically,
be combine the
perhaps necessary
to two
types.
hirta subtype
Euphorbia
Pollen grains without an operculum.
Euphorbia hirta Linn. Euphorbia hirta subtype; subprolate.

Soeprata 20 H [U] P = 30 E =
23 P:E =
1,30.
/x n
P.A.I. = 0,10.
Euphorbia peplus Linn. Euphorbia hirta subtype. Oblate
Fresh material, Utrecht. spheroidal.

P =
26,5 p E =
28,5 P:E =
0,93.
p
P.A.I. =
0,25.
Euphorbia palustris Linn. Euphorbia hirta subtype. Prolate
Fresh material, Hortus Utrecht. spheroidal.

P = 50 n E =
45 n P:E =
1,10.
P.A.I. = 0,3.
Diplocyathium capitata (Reichb.) Euphorbia hirta subtype; oblate spheroidal.
H. Schmidt P 31,5 fi E
= 34,5 fi P:E =
0,91. =
1891 P.A.I. 0,2.

Baldacci, Aug. [BR] =
Euphorbia esula subtype

Pollen grains with an operculum.
in the
This subtype differs from the other pollen grains Euphorbia
hirta type by its opercolum.
Euphorbia esula Linn. Euphorbia esula subtype. Oblate spheroidal.
Fresh material, Utrecht. P = 43 E = 40 P:E =
0,93.
n n
P.A.I. =
0,2.
Dichostema type
oblate Pollen in polar view circular to three-

Tricolporate; spheroidal. grains
lobed.
small absent.
Colpi narrow; margo or
reticulate. Columellae large. Exine thick.

Intectate;
The Dichostema type is differentiated from the other

types by its
distinct reticulum and narrow The columellae are fused

margo.
and form reticulum. The
laterally in their upper part a tigillate
have thick exine.
pollen grains a
Dichostema Pierre Dichostema oblate spheroidal-sub-

glaucescens type;
Klaine 1129 [L] PI. XXI, 4 oblate.
P = 31 /a E = 36 P;E = 0,87.
m:e=0,5. P.A.I.=0,3. Lumina 1-3 ft.
Anthostema aubryanum A. Juss. Dichostema oblate spheroidal.

de
type;
Spuitebanto 1074 [U] P = 29 E =

31,5 P:E =
0,92.
n n
Lumina 1-2 u.
Senefeldera macrophylla Ducke Dichostema

type; spheroidal,
For. Dept. 7549 [U] PI. XXI, 6 p = E = 36 P.A.I. =
0,2-0,25.
ft.
Lumina 2-3 fx.
Euphorbia cotinoides Miquel Dichostema prolate spheroidal.

type;
Ellenberg 1635 P 28 E 26 P:E 1,08.
[U] =
/i
=
n
=
Stahel and 816 P.A.I. ab-

Gongryp [U] m:e =
0,5. -
0,25. Margo
Lumina 2-3
sent; /x.
Stenadenium type
Tricolporate; subprolate. Pollen grains large.

Colpus transversal is broad and elongated; costae.
and
Colpi narrow long; costae
colpi indistinct. Margo absent.
Tectate; psilate; not reticulate. Columellae distinct but small in diameter.
Exine relatively thin.
Pollen than 40 and

grains of this type are large (larger /a) have
a relatively thin exine. The columellae are slender; their diameter is
small. In
polar view the pollen grains are not distinctly three-lobed.
A margo is absent.
The of Hura have shorter

pollen grains crepitans a colpus transversalis
than those of Stenadenium and Synadenium. The
shape and structure,
however, are similar.
Stenadenium Pax Stenadenium

spinescens type; subprolate.
Goetze 1099 [BM] PI. XXI, 1 P = 59 n E = 47 n P:E =
1,26.
Elaeophorbia drupifera (Thonn.) Staff Stenadenium
type.
Leeuwenberg 2314 [U] PI. XXI, 2 P =
58 fx E =
50 n P:E = 1,16.
P.A.I. =
0,2. Tectum perforatum.
Hura Linn. Stenadenium type.
crepitans
B. W. 694 [U] P = 75 n E = 63 P:E =
1,20.
fi
B.B.S. 502 [U] PI. XXII, 2 P.A.I. =

0,4.
Synadenium arborescens Boiss. Stenadenium
type.
D’Alleizette, Medley 8492 [L] Y
oung.
Pedilanthus
type
Tricolpate; oblate spheroidal to suboblate. Pollen grains slightly three-lobed.

Colpi broad. Margo narrow.
Membrana granulata.
Intectate (tectum perforatum?); psilate. Columellae tall.
Pedilanthus differs from other in the

types Hippomane configuration
the P.A.I. is
by the absence of a colpus transversalis. Also somewhat
larger.
Pedilanthus palmeri Millsp. Pedilanthus type.

Hinton 15793 PI. 1 P 72 E 83 P:E
[U] XXII, =
n
=
n
=
0,87.
P.A.I. =
0,3.
Pollen grains not placed in one of the Crotonoideae con-
figurations
Dalechampia type
Tricolporate; Pollen grains large, usually prolate to prolate spheroidal.

Colpus transversalis elongated. Sometimes costae quatoriales.
Colpi short and narrow. No costae colpi present.
Tectate; intra-reticulate. Reticulum coarse. Between the muri of the
very
reticulum a membrane is
present, covering the lumina.
102 W. PUNT
plate xxii.
1. Pedilanthus 2. Hura crepitans; 3. Actinostemon
palmeri;Ex- 4.
concolor;
coecaria bicolor.
The type is characterised by its large pollen grains and wide lumened
reticulum. The pollen grains that other type be
are so
typical, no can
them.
compared with
Dalechampia is easily distinguished from other genera by its typical

involucre. For this reason
Mueller
(1866) placed it in the vicinity
of the Bentham the similar characters
Euphorbieae. (1880) pointed to
in the Plukenetieae : e.g. climbing growth, stinging hairs and thickened

in the female flowers. Pax and K. Hoffmann
style (1919) thought
it better the into tribe. Croizat
to
separate genus an
apart (1940a)
~
and Hurusawa (1954) the genus

once
again placed near
Euphorbia
and its allies.
Pollen show and neither

grains a
typical structure are
comparable
with the in the Plukenetinae with those in the
types nor Euphorbieae.
Pax and K. Hoffmann’s classification maintaining the genus
in
a tribus not too far from the Plukenetinae the best

separate seems
opinion.
Dalechampia dioscoreifolia Poepp. et Dalechampia type; prolate.

Endlich. P 105 E 70 P:E
=
n
=
n = 1,50.
G. Klug 4186 [U] PI. XXIII, 6 Lumina to 12 Costae equatoriales.
up ft.
Dalechampia affinis Muell. Arg. Dalechampia type; prolate spheroidal.

B. W. 2183 [U] P = 105 E =
97 P;E =
1,08.
fi fi
Costae equatoriales.
Dalechampia brasiliensis Lam. Dalechampia prolate spheroidal.
type;
Y. Mexia 5270 [U] P = 57 E = 55 P:E =
1,04.
fi n
scandens Linn. Dalechampia type; oblate spheroidal.
Dalechampia
Serv. Forest. 4328 [U] P = 105 E = 75 P:E = 1,40.
n n
Dalechampia spathulata (Scheidw.) Baill. Dalechampia type; oblate spheroidal.
Fresh material, Hort. Utrecht. P = 6‘3yu E=65|< P:E =
0,97.
PI. XXIII, 5 Costae transversales; m:e =
0,5.
Hamilcoa
type
Tricolporate; suboblate. In polar view the pollen grains circular to slightly

convex triangular.
Colpus transversalis narrow.
Colpi narrow and short; indistinct costae colpi.

Tectate intra-reticulum fine not reticulate; columellaedistinct but small.
(?); or
The Hamilcoa be in of the discussed

type cannot
placed one con-
figurations. The pollen grains have columellae with small but distinct
capita and are for that reason
excepted from the Mallotus configura-
tion The of Hamilcoa would be in better agreement
(p. 77). structure
with the
Claoxylon configuration, (p. 90), but the related pollen grains
of and Pimeleodendron reticulate; the
Plagiostyles are moreover, polar
view differs. Perhaps the Moultonianthus in the Bernardia con-
type
is related the Hamilcoa of its
figuration (p. 73) type because
most to
circular polar view and fine reticulum in

present some species.
As the position of this type is still uncertain the author prefers
it from all other
to keep apart configurations.
104 W. PUNT
plate xxiii. I. Plagiostyles africana; 2. Pimeleodendron zoanthogyne; 3. Spirostachys

venenifera; 4. Hamilcoa zenkeri; 5. Dalechampia spathulata; 6. Dalechampia dioscoreifolia.
Prain
(1913) thought Hamilcoa was nearest related to Plagiostyles
Pierre and Pimeleodendron Hasskarl. Pax and K. Hoffmann
(1912b,
with the of and
1931) agreed relationship Plagiostyles Pimeleodendron,
which both in the
they placed Hippomaneae. Hamilcoa, .......
however, was
transferred to the Gelonieae. In doing so they rejected the affinity

between Hamilcoa and
Plagiostyles.
The three have several characters in common:
genera
1. Disc absent in male as

well as
in female flowers.
2. In male flowers an ovarium rudiment is absent. Filaments are
very
short or wanting.
3. Pedicels are thickened.
4. undivided.
Styles are
5. Plants dioecious (In Hippomaneae usually monoecious).
Perhaps Nealchornea, in the placed in the Sumbavia

present paper
is also related Hamilcoa and its allies.

configuration (p. 69), to
Hamilcoa zenkeri (Pax) Prain Hamilcoa suboblate to oblate

type;
Zenker 4765 [L] PI. XXIII, 4 spheroidal.
P =
23,5 E = 27 P:E =
0,87.
n
m: e < 0,5. P. A.I.> 0,5. Lumina ca. 1

/i.
Pimeleodendron zoanthogyne J. J. Smith Hamilcoa spheroidal.

type;
Hallier 152a [U] PI. XXIII, 2 P = E = 24 m:e ca. 1. P.A.I. =
fi.
Plagiostyles africana (Muell. Arg.) Prain Hamilcoa type; oblate spheroidal.

Zenker 94 PI. XXIII, I P=27 E 29yit P:E 0,90.
[U] /*
= =
m:eca. 1. P.A.I. =
0,4. Costae colpi.
Taxonomic comment on the Crotonoideae
The subfamily of the Crotonoideae includes a large number of pollen

types.
The types with a attract most attention. When in
croton-pattern
the future revision of the Crotonoideae will be the
a undertaken,
placing of all genera with a
croton-pattern into one group will be
advisable. This will benefit a natural classification of the subfamily.

The of, with Cunuria and Garcia with
relationship e.g., Joannesia
Sagotia will, in this way,
show to better advantage than it does at
present in Pax and K. Hoffmann’s system.

Two other of the Cnesmosa
groups well-distinguished pollen grains,
Plukenetia found in the sub-
configuration and the configuration, are
tribe Plukenetiinae. Omphalea is added to the Plukenetia configuration,

which with the opinion of Croizat 62).
agrees (p.
Mueller Arg., Bentham and Pax and K. Hoffmann include the
tribe Chrozophoreae in their This tribe falls into two

systems. groups
of The first with e.g. Aleurites,
pollen types. group Crotonogyne, etc.
possesses pollen grains with a croton-pattern. As stated previously

these should be from the other and transferred
genera separated ones
to the group of which Croton is the principal genus. The second group
with, e.g., Sumbavia, Chrozophora, Agrostistachys, has aperturate pollen

106 W. PUNT
which belong several pollen The author

grains, to
types. present
would like retain this second group the subtribe
to as Chrozophorinae.
The tribe of Pax and K. Hoffmann includes
Acalypheae most
of the Crotonoideae. There are, however, comparatively few

genera
which related ( Mallotus configuration,
types are, moreover, closely
Bernardia configuration, etc.).
The tribes Gelonieae and Cluytieae, too, include genera of which the
pollen grains are provided with a croton-pattern mixed with

up
of which the pollen grains lack this structure. These latter
genera
are divisible into several A reclassification of these

genera types.
tribes is
badly needed.
Although Hippomaneae and Euphorbieae are both homogeneous taxa
which are
taxonomically well-defined, the
morphology of the pollen
shows but little difference between the Pax (1924)
grains two
groups.
the these groups

already pointed out relationship of on other grounds.
Since the Hippomaneae are treated by all authors as a tribe of the
Crotonoideae, the Euphorbieae should also be treated as a tribe of the
Crotonoideae close to the Hippomaneae.
SUMMARY
In the study pollen morphology of the Euphorbeaceae is treated

present as an
additional character in
taxonomy.
Besides the of the occurring in the of Pax and K. Hoff-
greater part genera system
of the published after 1931 studied.
mann
(1931), most genera are
The pollen grains have been described with the aid of a terminology as simple
as possible. In principle the terminology of Iversen and Troels-Smith has been
followed, although in addition, improvements of Erdtman have been used.

many
One of the simplifications is the rejection of Potoni6’s term sculpture. All elements
the endexine called all elements

occurring on are structure elements; structure
together form the structure of a pollen grain. For the sake of endexine
consequence
and extexine discussed separately.

apertures apertures are
Different pollen grains are placed in different pollen types. If the differences are
of minor importance, the pollen grains are placed in subtypes. Several

types
can
have characters in To the correspondences, these

some common.
express types are
assembled in configurations.
As the pollen types in Phyllanthoideae and Crotonoideae differ distinctly, the division
of the Euphorbiaceae in these subfamilies is maintained m the discussion of the results.
The be separated in three of pollen

Phyllanthodieae can large groups types ( Antidesma
Amanoa and which
configuration, configuration Aristogeitonia configuration), agrees
with the grouping of Pax in 1924. The remaining small configurations belong in
taxonomic to the of the Antidesma configuration.

respect genera
In the Crotonoideae pollen grains with

many genera possess a
croton-pattern.
These genera should be treated as a single group.
Besides this natural the Plukenetiinae possess pollen grains which are clearly
group,
distinguished from other in the Crotonoideae. Pollen grains of Omphalea are

genera
similar to those in the Plukenetia configuration. This pollen-morphological result
with the opinion of Croizat.

agrees
The remaining pollen grains in the Crotonoideae are less to differentiate in

easy
One of the configurations is the Mallotus configuration, which

groups. largest
includes most of the Acalypheae and several or other tribes. The
genera genera
Hippomane configuration is another large one. This configuration comprises the
tribes The of both tribes

Hippomaneae and Euphorbieae. pollen grains are
very
similar. The Pachyslroma is pollen-morphologically as well as taxonomically

genus
related to the tribe Hippomaneae.
Per a. treated Pax K. is related its

as a
separate tribe by and Hoffmann, by pollen
grains to some
genera
in the Acalypheae.
Dalechampia is habitually related to the in the Plukenetiinae. Pollen-

genera
morphological data, however, do not this relation. The pollen grains of

support
not similar to other pollen

Dalechampia are
any type.
Stenolobeae is in the
The morphology of the pollen grains of the
agreement with
opinion of Pax, that separation of these Australian is an artificial one.

any genera
REFERENCES
H. K. 1949. Note the Euphorbiaceous Botryophora. Kew

Airy-Shaw, on
genus
Bull. 3: 484.
1960. Notes on Malaysian Euphorbiaceae. Kew Bull. 14: 353-397,

.
469-475.
Baillon, H. 1858. Etude du des Paris. 684

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Bot. Mus. Leaf!. Harv. Univ. 12: 325-349.
Bentham, G. 1880. Notes on Euphorbiaceae. Journ. Linn. Soc. Bot. 17: 185-267.
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C. L. 1825-1826. Bijdragen de flora Ned. Ind. Batavia. 1169
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Arb. 21: 76-107.
—
1940c. New and critical Euphorbiaceae from Eastern Tropical Asia.
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Journ. Arnold Arb. 21: 490-510.
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1941a. The tribe Plukenetiinae of the Euphorbiaceae in Eastern Tropical
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.
ser. 3. 17: 204-208.
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1942a. Notes on the Euphorbiaceae 3. Bull. Bot. Gard. Buitenzorg
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1943a. Notes on American Euphorbiaceae. Journ. Wash. Acad. Sci.
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1943b. New and critical Euphorbiaceae of Brazil. Tropical Woods 76:
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.
1943c. Novelties in American Euphorbiaceae. Journ. Arnold Arb. 24:
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1945. New or
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1946a. Nomenclatoral transfers and corrections in the Euphorbiaceae.

.
Tropical Woods 88 : 30-32.

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1946b. Novelties in Journ.
289-291.
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INDEX
(Asterisks indicate illustrations)
*
Acalypha type, 79, 81 xiphoclada, 95
Acalypha aronioides, 81 Amyrea humberti, 86
diversifolia, 81 Anabaena, 10
�
indica, 81 10
Anabaenella,
racemosa, 81 tamnoides, 63
platyphylla, 81 10
Anabaina,
*
scandens, 81 Andrachne subtype, 28
6
Acalyphopsis, Andrachne
aspera, 29
Acidocroton adelioides, 51 colchica, 29
*
Acidoton urens, 57 phyllanthoides, 29
*
Acephila excelsa, 29, 32 telephioides. 29
*
Actephilopsis malayana, 60 Androstachys johnsonii, 43
.
,
‘
�
Actinostemon concolor, 97
Angostyles longifolia, 63
lanceolatus, 97 Annesijoa, 6
Adelia ricinella, 68 Anomalocalyx uleanus, 51
Adenochlaena leucocephala,
. 94 Anthostema aubryanum, 101
Adenocline subtype, 48, 54 Antidesma configuration, 20, 30, 32,

Adonocline, 48, 54 34, 39, 36, 43, 46
56 Antidesma 19, 20, 21, 26, 28

acuta, type,
*
mercurialis, 56 24
Antidesma subtype, 20, 21, 22,
Adenopeltis colliguaya, 99 Antidesma 22
agusanense,
Adenophaedra, 73 *
bunius, 22
minor, 75, 82 diandrum, 22
prealtum, 82 edule, 22
Adriana, 96 ghesaembilla, 22
quadripartita, 80 22
membranaceum,
Agrostistachys, 105 obovatum, 22
indica, 76 22
venosum,
*
sessilifolia, 76 Aonikena 66
patagonica,
*
Agyneia bacciformis, 39 Aparisthmium cordatum, 83
Alchornea 85
type, 19, 73, 79, 82, 83, Apodandra, 62
Alchornea, 73, 75, 85, 94 buchtienii, 63
ilicifolia, 83 Apodiscus chevalieri, 22

laxiflora, 82 Aporosa, 25
82 dioica, 24
rugosa,
schomburgkii, 82 frutescens, 24
*
82 lindleyana, 24
triplinervia,
*
villosa, 82 Aporosella, 25
*
Alchorneopsis, 85 Arachnodes, 38
floribunda, 86 chevalieri, 40
86 68
trimera, Argythamnia type,
Aleurites, 105 Argythamnia, 66
moluccana, 50 *
68
candicans, 266,
Algernonia brasiliensis, 99 *
Argomuellera macrophylla, 76
Alphandia furfuracea,.50 Aristogeitonia configuration, 40, 46
Amanoa configuration, 32, 36, 38, 46 Aristogeitonia 40

type,
Amanoa 32, 33, 34 41
type, Aristogeitonia subtype,
Amanoa, 19, 32, 33 Aristogeitonia limoniifolia, 41
boiviniana, 33
Astrocasia, 25, 32
bracteosa, 33 tremula, 26
grandiflora, 33
Astrococcus, 62
* *
33
guianensis, cornutus, 63
�
oblongifolia, 33 Athroandra 92
africana,
*
Amperea type, 19, 94 mannii, 92
Amperea spartioides, 95 Avellanita bustillosii, 80
Baccaurea subtype, 21, 22, 24 Cenesmon, 10, 57

Baccaurea javanica, 24 Centroplacus glaucinus, 89
24 Cephalocroton, 96
oxyxarpa,
racemosa, 24 cordofanus, 94
* *
sumatrana, 24 94
Cephalocrolonopsis socotrana, ,
Baliospermum, 48 Cephalomappa configuration, 68
montanum, 51 Cephalomappa type, 68
Baloghia lucida, 51 Cephalomappa malloticarpa, 69

Chaetocarpus subtype, 90
Benoistia, 48
*
perrieri, 51 Chaetocarpus castanocarpus, 90
Bemardia configuration, 73, 92, 103, schomburgkianus, 90
106 Cheilosa 79, 90

type,
*
Bernardia 73, 75, 76, 77 Cheilosa montana, 90
type,
Bernardia, 85 Chiropetalum configuration, 62, 65
corensis, 75 Chiropetalum type, 65, 66, 68
lorentzii, 75 Chiropetalum 66
canescens,
* *
pulchella, 75 lanceolatum, 66
Bertya, 48 66
patagonica,
gummifera, 53 Chlamydojatropha, 6
Beyeria, 48 Chondrostylis bancana, 85

leschenaultii, 53 Chonocentrum, 21, 33
Bischofia 43 cyathoforum, 25
type,
Bischofia, 9, 40, 44
Chrozophora type, 69, 70, 72
�
javanica, 44 Chrozophora, 105
Blachia umbellata, 51 *
70
plicata,
Blotia oblongifolia, 34 70
tinctoria,
Blumeodendron 79, 81 Cladogelonium, 48, 54
type,
*
Blumeodendron tokbrai, 81 *
54
madagascariense,
Bocquillonia sessiliflora, 83 configuration, 69,
Cladogynos 93
Bonania cubana, 99 Cladogynos type, 69, 93, 94
domingensis, 99
Cladogynos
*
Botryophora geniculata, 81 orientalis, 94
Breynia configuration, 38 Claoxylon configuration, 90, 103
Breynia 38, 39 Claoxylon type, 92, 93

type,
Breynia subtype, 38 *
Claoxylon cuneatum, 92
Breynia, 29, 38, 39, 44 tumidum, 92
*
fruticosa, 39
Claoxylopsis, 5
nivosa, 39 Clarorivinia, 6
39 Cleidion, 82, 83
racemosa,
Breyniopsis subtype, 39 80
angustifolium,
Pierrei, 39
* *
Breyniopsis javanicum, 80
Briedelia 32, 36 Cleistanthus, 32

type,
Briedelia, 9, 32 dichotomus, 34
*
assamica, 36 ferruginous, 36
glauca, 36 34
winkleri,
*
monoica, 36 Clutia type, 19, 75, 77
36 Clutia abyssinica, 77
stipularis,
alaternoides, 77
Calpigyne hainanense, 86 natalensis, 77
*
Calicopeplus, 5 paxii, 77
Caperonia type, 72 rubricaulis, 77
Caperonia subtype, 72 Cluytiandra madagascariensis, 34
Caperonia, 69 Cnesmosa configuration, 57, 58, 105
corchoroides, 72 Cnesmosa 57, 58

type,
*
palustris, 72 Cnesmosa, 10
serrata, 72 javanica, 57
Carumbium, 10 Cnidiscolus stimulosis, 53
Caryodendron type, 72 urens, 53
Caryodendron grandifolius, 73 Coccoceras borneënse, 80

�
orinocense, 73 *
Codiaeum variegatum, 51
Cavacoa aurea, 51 stellingianum, 51

112 W. PUNT
*
Coelebogyne ilicifolia, 83 Ditaxis catamarcensis, 66
Coelodepas, 10 diversiflora, 66
Coelodiscus lanceolatus, 80 fasciculata, 66

*
Colliguaya integerrima, 97 fendleri, 66
Cometia, 5 lancifolia, 66
Conveiba lutifolia, 83 salina, 66
simulata, 83 Ditta, 5
Conceveibastrum martianum, 83 Dodecastigma, 48
Cordemoya integrifolia, 80 amazonicum, 53
Corythea, 5 *
Domohinea perrieri, 53
Croizatia, 5 Drypetes, 21, 44

*
Crotonoideae, 47 glauca, 28
Croton 47, 58, 90 laterifolia, 28
configuration,
Croton type, 48, 53, 54, 57, 58 macrophylla, 28
Croton bahamensis, 50 variabilis, 28
*
cuneatus, 50 Duvigneaudia inopinata, 99
hirtus, 50 Dysopsis configuration, 60
lingiradiatus, 50 Dysopsis 60
type,
* *
matourensis, 47, 50 Dysopsis glechomoides, 60
pullei, 50
*
Crotonogyne, 105 Elaeophorbia drupifera, 101
parvifolia, 50 Elateriospermum, 48
preussii, 50 51
tapos,
*
Crotonogynopsis usambarica, 85 Eleuterostigma lehmannianum, 62
50 48
Crotonopsis elliptica, Endospermum,
Cunuria, 49 moluccanum, 54
bracteosa, 50, 53 Epiprinus poilanei, 94

spruceana var.
Cyathogyne spathulifolia, 22 Eremocarpus, 48
viridis, 22 setigerus, 50
Cyrtogonone argentea, 50 Erismanthus 77

type,
*
Cyttaranthus 76 Erismanthus indochinensis, 77
congolensis, 75,
*
oblique, 77
Dalechampia 101 Erythrococca africana, 92
type,
Dalechampia affinis, 103 mannii, 92
brasiliensis, 103 stolziana, 92
*
dioscoriifolia, 103 Euphorbia hirta 96, 100
type,
scandens, 103 Euphorbia esula subtype, 100
*
spathulata, 103 hirta 100
Euphorbia subtype,
Dalembertia populifolia, 99 Euphorbia, 9, 103
30 101
Danguyodrypetes ambigua, cotinoides,
Dendrophyllanthus type, 37 esula, 100

�
Dendrophyllanthus ripicolus, 37 hirta, 100
Deuteromallotus acuminatus, 80 palustris, 100
Deutzianthus tonkinensis, 50 peplus, 100
*
Dichostema 96, 100 Excoecaria bicolor, 97
type,
*
Dichostema glaucescens, 100 phillippinensis, 99
Dicoelia 20, 21, 28, 29, 32, 44 venenifera, 97

type,
Dicoelia subtype, 28
Dicoelia affinis, 28 Flueggia, 25

Dimorphocalyx murina, 51 Fontainea pancheri, 51
Diplocyathium capitata, 100 Fragariopsis, 62

*
Discocarpus 32, 36 scandens, 63
type,
Discocarpus, 21
*
essequeboensis, 36 Galearia dognaiensis, 89
34 filiformis, 89
hirtus,
Discoclaoxylon hexandrum, 92 Garcia, 48, 105
Discocleidion 79, 83 50
type, nutans,
»
Discocleidion rufescens, 85 Gatnaia annamica, 24
Discoglypremna caloneura, 76 Gavarretia terminalis, 83

*
Dissiliaria tricornis, 41 Gitaria, 6
Ditaxis 65, 66, 68 Givotia rottleriformis, 51

type,
Glochidion 39 longifolium, 86
type,
Glochidion, 29, 38, 44 *
Kunstlerodendron sublanceolatum, 85
concolor, 39
littorale, 39 Lachnostylis hirta, 34
obscurum, 39 Lasiochlamys, 5
ramiflorum, 39 Lasiococca symphylliaefolia, 86

*
sericeum, 39 Lasiocroton 65, 68
type,
*
superbum, 39 Lasiocroton macrophyllus, 68
Glycydendron amazonicum, 54 Lautembergia coriacea, 83
Glyphostylus laoticus, 97 multispicata, 83
Godefroya rotundata, 36 Leidesia 79, 88

type,
Grimmeodendron eglandulosum, 99 Leidesia, 54, 60
*
Grossera major, 50 procumbens, 88
Gymnanthes lucida, 97 Lepidoturus laxiflorus, 82
jamaicensis, 97 Leptonema type, 29
*
Leptonema venosa, 29
*
Haematostemon guianensis, 63 Leucocroton flavicans, 66
�
Hamilcoa 103 Lingelsheimia frutescens, 30
type,
*
Hamilcoa zenkeri, 105 Longetia buxoides subtype, 41
* �
Heterocalyx laoticus, 76 Longetia buxoides, 41
Hevea 56 carunculata, 41
type, 48,
*
Hevea brasiliensis, 56 gynotricha, 43
guianensis, 56 malayana, 41
Heywoodia type, 20, 30

*
30 Mabea 97
Heywoodia lucens, caudata,
Hippomane configuration, 95 fistulifera, 97
Hippomane type, 100 97
19, 96, indorum,
Hippomane subtype, 96 piriri, 96
*
Hippomane mancinella, 96 taquari, 96
Holstia tenuifolia, 50 Macaranga densiflora, 80
Homalanthus, 10 harveyana, 80
�
*
Homonoia 86 spinosa, 80
javensis,
riparia, 86 Maesobotrya dusenii, 22
Hura, 96 floribunda, 22
crepitans, 101 Mallotus configuration, 77, 90, 92,

*
Hymenocardia type, 44 103, 106
Hymenocardia, 21, 44 Mallotus 85, 92

type, 19, 79, 81, 82,
acida, 46 Mallotus subtype, 79, 80
ulmoides, 46 Mallotus, 70, 79, 94
* �
Hyaenanche globosa, 41 albus, 80
Hyeronima, 21 claoxyloides, 80
alchorneoides, 22 miquelianus, 80
*
laxiflora, 22 repandus, 80
Manihot 47, 48, 53, 54

type,
Jatropha, 53 Manihot, 53, 96
integerrima, 51 esculenta, 53
multifida, 51 saxicola, 47, 53

�
3 Manniophyton fulvum, 50
panduraefolia,
Joannesia, 48, 105 Maprounea brasiliensis, 99
princeps, 51 guyanensis, 99
Julocroton argenteus, 50 Mareya type, 79, 83, 86, 92

�
triqueter, 50 Mareya brevipes, 85
Mareyopsis longifolia, 85
*
Keyodendron, 5 Margaritaria nobilis, 26
Klaineanthus 47, 48, 54, 56 Martretia 46
type, type,
*
Klaineanthus subtype, 54 Martretia quadricornis, 46
*
Klaineanthus, 48, 54, 56 Megistostigma peltatus, 57
* �
gaboniae, 54 malaccense, 57
�
Koilodepas, 10 Melanolepis multiglandulosa, 70
*
86 Mercurialis 93
bantamense, type,
hainanense, 86 Mercurialis, 88
114 W. PUNT
*
93 41
annua, Paragelonium perrieri,
reverchonii, 93 Pausandra densiflora, 51
93 morisiana, 51
tomentosa,
Mettenia, 5 Pedilanthus 101

type,
*
Micrandra, 47, 48, 50 Pedilanthus palmeri, 101
brownsbergensis, 50, 54 Pentabrachion 34

o ,
reticulatum,
� .
siphonioides, 50, 54 Pera 79, 86, 88
type,
*
Micrantheum ericoides, 43 Pera bicolor, 88
hexandrum, 43 glabrata, 88
Micrococca mercurialis, 92 Petalostigma quadriloculare, 41
Microdesmis 79, 88 Philyra subtype, 69, 72

type,
*
Microdesmis subtype, 89 Philyra brasiliensis, 72
*
Microdesmis caseariaefolia, 89 Phyllanoa, 5
paniculata, 89 Phyllanthoideae, 20
zenkeri, 89 Phyllanthus nutans configuration, 37

Mildbraedia paniculata, 51 Phyllanthus acuminatus type, 43
Mischodon zeylanicus, 41 Phyllanthus 37
nutans type,
Monadenium, 5 Phyllanthus acidus subtype, 20, 26

*
Monotaxis grandiflora, 70 Phyllanthus niruri subtype, 24
.
„ .
Moultonianthus
type, 19, 76, 103 Phyllanthus pentaphyllus subtype,
Moultonianthus, 73, 75, 76 21, 24, 33, 77
*
leembruggianus, 77 Phyllanthus, 18, 20, 21, 25, 29, 38, 77
acidus,, 25, 26
�
Nealchornea 73
type, acuminatus, 43
Nealchornea, 105 acuminatissima, 26
*
yapurensis, 73 adianthoides, 37
Necepsia afzelii, 86 24
amarus,
*
Neoboutonia, 47, 48 capillaris, 26
macrocalyx, 51 discoideus, 26
*
Neomphalea, 6 elsiae, 25, 26
Neopalissya castaneifolia, 85 guianensis, 25
Neoroepera banksii, 43 hyssopifolioides, 25
*
Neoscortechinia arborea, 90 muellerianus, 26
*
Neotrewia cumingii, 83 niruri, 25
*
Neotrigonostemon, 6 37
nutans,
*
Nephrostylus poilanei, 86 pentaphyllus, 24
reticulatus, 26
Oldfieldia, 44 stipulatus, 24
africana, 41 sublanatus, 24
Oligoceras, 48 speciosus, 37
eberhardtii, 50 urinaria 25
*
Omalanthus nutans subtype, 19. 96, Pimeleodendron zoanthogyne, 105
97 Piranhea, 44
*
Omalanthus, 10 trifoliata, 41
* *
97 Plagiostyles africana, 105
nutans,
populneus, 96 Platygyne 57, 58
type,
*
Omphalea, 62, 105 Platygyne hexandra, 58
*
diandra, 63 Pleiostemon 30
type,
*
macrophyllum, 99 Pleiostemon 30
Ophthalmoblapton verrucosus,
Ostodes paniculata, 51 Plukenetia configuration, 60, 65, 105
pendulus, 51 Plukenetia 19, 62

type,
Plukenetia verrucosa sybtype, 63
Pachystroma 99 Plukenetia volubilis subtype, 62

type,
Pachystroma, 96 Plukenetia, 62
*
longifolium, 100 abutifolia, 62
Pachystilidium 65 buchtienii, 63
type,
hirsutum, 65 conophorum, 63
Pachystylidium
Paivaeusa dactylophylla, 41 corniculatus, 63
Pantadenia, 48 tamnoides, 63
*
adenanthera, 51 verrucosa, 63
*
Paradrypetes, 5 volubilis, 62
*
Pogonophora schomburgkiana, 90 virosa, 26
Polyandra, 82, 83 Seidelia type, 79, 88
Poranthera corymbosa, 29 Seidelia, 54, 60
microphylla, 29 triandra, 88
*
Prosartema gaudichaudii, 60 Senefeldera macrophylla, 105
*
Protomegabaria macrophylla, 22 Senefelderopsis, 6
*
Pseudagrostistachys africana, 76 Speranskia type, 73, 75, 95
*
Pseudanthus nematophorus, 43 Speranskia pekinensis, 76
*
pimeleoides, 43 Sphaerostylis, 57
Pseudocroton 95 63
type, natalensis,
Pseudocroton tinctorius, 95 tulasneana, 57, 63
*
Pseudolachnostylus glauca, 34 Sphyranthera, 6
*
Pterococcus corniculatus, 63 Spirostachys venenifera, 97
Ptychopyxis 81 Spondianthus preussii, 22

costata,
kingii, 81 Stachystemon 41
type,
*
Putranjiva 44 Stachystemon vermicularis, 43
type,
*
Putranjiva roxburghii, 44 Stenadenium 101
type,
*
Pycnocoma subtype, 80 Stenadenium spinescens, 101
*
Pycnocoma 80 Stillingia 97
cornuta, gymnogyna,
macrophylla, 81 Strophioblachia fimbricalyx, 51
Sumbavia configuration, 69, 73, 105
Ramelia, 6 Sumbavia 19, 69, 70, 72

type,
Reverchonia, 21 Sumbavia, 105
* *
arenaria, 24 rottleroides, 70
Richeria 26 Sumbaviopsis albicans, 70

type,
Richeria laurifolia, 28 Suregada, 48, 53
Richeriella, 21 glomerulata, 53
*
Richeriella gracilis, 26 subglomerulata, 53
Ricinocarpus, 48 Symphyllia siletiana, 94
pinifolius, 53 Synadenium arborescens, 101
Ricinodendron heudelotii, 51 Syndyophyllum, 6
Ricinus 19, 79, 86, 88

type,
* *
Ricinus communis, 86 Tannodia cordifolia, 53
Riseleya, 6 Tetracarpidium conophorum, 63
Romanoa, 10, 62, 63 Tetracoccus ilicifolius, 41
* 99
Tetraplandra gibbosa,
Sagotia, 48, 105 Thecacoris leptobotrya, 22
51 Tetrorchidium type, 56
racemosa, 48,
Sandwithia guyanensis, 53 Tetrorchidium, 48, 56
*
Sapium aubletianum, 99 didymostemon, 57
ellipticum, 99 rubrivenium, 57
klotzschianum, 99 Thelypetalum, 6
longifolium, 97 Thymelaeaceae, 19
*
99 Thyrsanthera suborbicularis, 70
montanum,
99 fallax 57
sebiferum, Tragia type, 19,
Sauropus androgynis, 39 Tragia tristis subtype, 63
Savia 32, 34, 36 Tragia, 18, 58

type,
Savia andringitrana, 34 capensis, 63
*
erythroxyloides, 24, 33 58
fallax,
sessiliflora, 34 geraniifolia, 65
Sebastiania chamaelea, 97 ramosa, 65
corniculata, 97 sellowiana, 58
*
schottiana, 97 stolziana, 63
*
19, 20, 21, 25, 26, tristis, 63
Securinega type,
32, 39 volubilis, 65
Securinega subtype, 25, 26 57

Tragiella,
*
Securinega, 25, 30, 32 Trewia nudiflora, 80
congesta, 22 Trigonopleura, 73
*
neopeltandra, 26 malayana, 90
26 Trigonostemon redioides 58
ramiflora, type,
suffruticosa, 25 Trigonostemon 58
verrucosus
type,
116 W. PUNT
Trigonostemon, 73 kirkiana, 36
fungii, 58
longifolius, 60 Vaupesia cataractarum, 53
*
rediodes, 60 Veconcibea 83
latifolia,
*
verrucosus, 58
Tritaxis, 48 Wetria macrophylla, 80
gaudichaudii, 51 Wielandia elegans, 34
Uapaca, 32 Zimmermannia 20, 29

type,
* *
heudelotii, 36 Zimmermannia capillepes, 29

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Wentia 7 (1-116)

Pollen morphology of the

euphorbiaceae with special reference

(received December 28th, 1961)

Chapter I General Introduction 2

Chapter III Material 5

c. Preservation of pollen grains 7

Chapter V Some nomenclatural remarks 9

Chapter VI Pollen morphology 10

Chapter VII Glossary 15

Chapter VIII Results 18

A. Pollen grains of the Euphorbiaceae 18

B. Discussion of the results

Phyllanthus nutans configuration 37

have stated Lindau Wodehouse

In the work the therefore extended

1931, The taxonomic conclusions, however, have

character. Future investigators of the

The have been carried the “Botanisch Museum

Herbarium”, Utrecht. The writer is indebted the

director, Professor Dr. for him the

the his institute and for his and

very grateful to the “Miquelfonds” which covered the expenses

The study of pollen morphology started in 1675, when Malpighi

observed, that various types of pollen grains are to be found in different

He variations of colour and Grew

smooth and spiny pollen grains. In the centuries

number of botanists enriched our knowledge of pollen morphology.;

that the of the the

features. Some later Fritzsche

published his principal work on pollen grains (Ueber den Pollen,

rately analysed. His drawing and description of Jatropha panduraefolia

accurately reproduces the construction of the (see

Pollen of related similar.

3: Sometimes unrelated have similar

In 1883 Radlkofer made of obtain

better classification of the Acanthaceae. It \yas done more

ganz streng durchgefiihrt werden soil”. It is obvious that, with regard

of the end of the 19th

diagram; the primitive pollen analysis had resulted

Potoni£ in 1934 and Wodehouse in 1935 published works on

pollen morphology that formed the basis of our modern terminology.

Potonie as well as Wodehouse needed a of well defined terms

excelled in exactitude and

made it determine which in

minoid? Nevertheless, his book is of valuable data

extensive notes on his own system.

Pfianzenreich”. In the second edition of Engler and Die

made on the system of the Euphorbiaceae. Most of them were given in

several Schultes discussed the Hevea and its

genus Phyllanthus into subgenera and sections.

Herbarium material obtained from the herbaria

used in the study;

Kew (K); The Herbarium, Royal Botanic Gardens.

Leiden (L): Rijksherbarium.

Wageningen (WAG): Laboratory for Plant Taxonomy and Plant Geography.

the of has been avoided. in

has been used for

possible all known genera of the Euphorbiaceae were examined. In a

number of genera it for various reasons,

1. Male flowers unknown.

2. The material studied did contain male flowers,

few, or unripe ones.

Cometia Thouars ex Baillon