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Pollen morphology and taxonomy of
section Cera-
manthus baillon S.L. of the genus Phyllanthus
(Euphorbiaceae)
W. Punt
and
Laboratory ofPalaeobotany
Utrecht
Palynology, Botanical
Museum
and
Herbarium,
State
University,
(The Netherlands)
November
(Received
26, 1971)
ABSTRACT
Punt, W.,
1972. Pollen
morphology and
taxonomy
of section
Three sections
The
with
a
total number of four
pollen grains show
a
strong
resemblance
differentiate between the three sections
taxonomic
Mueller
reasons
have
Arg.
the sections
been united
proved
Ceramanthus
into one
of the
species
to each
Ceramanthus Baillon
13:
Phyllanthus (Euphorbiaceae). Rev. Palaeobot. Palynol.,
genus Phyllanthus
other and
to be
rather weak.
viz. section
have
also the taxonomic
Baillon, Cluytiopsis
section;
s.l. of the
genus
213-228.
Because
Mueller
Ceramanthus
of both
Arg.
been
examined.
arguments
to
palynological
and
and Anisolobium
Baillon
s.l.
INTRODUCTION
In
a
recent
albodiscus
Hutch, and P.
Pax
regard
variety
only conspecific
species
Pax
Shaw made the
newly proposed
by
pollen morphology
occurred
to
appeared
problems
mentioned by
Airy
same
was
(1956).
shows
large diversity
in this
paper he
P.grahamii
Africa,
a
special
(1969)
published
P. beillei
some
and, especially,
species
Hutchinson
but also with
Ridley
and
rather unfortunate
unexpected
the last
P.stolzianus
et
M.B. Moss
an
so
East Asian
Phyllanthus
soon
in the genus Phyllanthus
pollen grains might
group of Phyllanthus
were
as
in Thailand and Cambodia. Besides this
Webster
Shaw
circle of affinity
that Cleistanthus albodiscus
which
that Phyllanthus
possible,
that the East African
my mind that the features of the
the taxonomic
it
same
P.albodiscus and P.beillei
K.Hoffm. and
conspecific,
are
the
In the
Sprengel occurring
discovery,
name
et
Arg.
to
with P.beillei from West
of P.cochinchinensis
ceramanthus Webster
taxa
the
(1969) thought
might belong
is of remarkable interest. It
were not
As
to
K.Hoffm., P.nyassae
et
Airy
Shaw
welwitschianus Mueller
discoveries with
species
Shaw
publication Airy
(Ridley) Airy
studied.
be of
species.
after the
(Punt, 1967),
help
to
For this
it
untangle
reason
several
214
W. PUNT
MATERIAL
The material
Bruxelles
The
taken from sheets which
was
(Belgium):
Herbarium and
(The Netherlands):
a
Botanical Museum and
Plant
present in the following herbaria: BR,
(Great Britain): Royal
and Plant
that the P.beillei material from Asia does
pity
of the Asian
Gardens,
Utrecht
Rijksherbarium; U,
So that it
Geography.
not
contain male flowers
possible
was not
with those of the African
specimens
Botanic
Herbarium; WAG, Wageningen (The
Taxonomy
Shaw, personal communication, 1971).
grains
are
Kew
Library; L, Leiden (The Netherlands):
Netherlands): Laboratory for
It is
1’fetat; K,
Jardin de
(Airy
compare the
to
pollen
ones.
METHODS
Preparation
Pollen
grains
were
described
by
Leins
to
sections up
and mounted
prepared
described by Reitsma (1969).
(1968)
The
according
pollen grains
with the aid of
a
were
to
cut
the
method
acetolysis
according
to
as
the method
Leitz microtome which enables the
as
cutting
of
1 p thin.
Descriptions
Pollen
grains
were
apochr. 40/0.65;
used for the
examined with
100/1.32
descriptions.
slide. This number does
present in
relative
can
rather
a
only
as
and
microscope having objectives
Leitz Ortholux
The
terminology suggested by
Measurements have been carried
not
simple
a
90/1.40.
pretend
to
way averages
the influence of the
be of considerable influence
on
give
out at ten
Reitsma
pollen grains
reliable statistical values, but the
and limits.
They
method
preparing
the sizes of the
with the
is
per
figures
should be considered
together
(1970)
to
be
mounting
method
1970;
pollen grains (Reitsma,
1970).
Praglowski,
Photographs
The
as
a
light microscopical photographs
that mentioned under the
Leitz Orthomat. To increase the
has been used. The film was
Besides these
been taken
Breukelen,
were
taken with the
heading “Descriptions”.
a
contrast an
Kodak
the
Recordak,
Cambridge equipment
The Netherlands.
same
microscope equipment
microscope
was
combined with
interference green filter AL 546
lightmicroscopical photographs
using
This
(546 nm)
AHU-microfilm 5460 film.
a
few
scanning-electron
photographs
have
of the “Micro-struktuur Laboratorium” at
POLLEN
MORPHOLOGY AND TAXONOMY OF CERAMANTHUS
BAILLON S.L.
215
TAXONOMY
Species
cochinchinensis
Phyllanthus
P.cochinchinensis
Loureiro
species.
Cathetus
the
as
a
P.cochinchinensis.
was
species
Sprengel
as
a new
had to choose
complete
a
(1804)
several years before. The
is
plant
North of Vietnam and the South of China up
Phyllanthus
albodiscus
The second
representative
name
(Ridley) Airy
Ceramanthus gracile,
Baillon
(1858), however,
a new
the latter
under the
name
necessary.
species
is
According
that
species
taxa
was
originally
(1927)
names
grahamii
of P. beillei. At the
Beille
and
(1969)
the
same
Phyllanthus
had
by
in the
under the
(1844),
of
too
already published
a new
specific
correct name must
Bangka, Malaya,
stolzianus Pax et K.
Hutchinson
time he
et
brought
M. B.
up
to
Rhodesia in the
South)
in
little value
to
another
name
to
taxon
taxon
was
be P. albodiscus.
The
the South of Thailand.
light
West- and East Africa
that this
to
Shaw
nyassae
(1969)
Pax
be considered
taxon
was
et
as
discovered
K.
synonyms
obviously
con-
as
not
a
of P.
variety
unique,
that
fasciculatus (var.
a taxon
(Tanzania, Kenya
can
species
to
the South of
Arg.
from this group, P. welwitschianus,
is
more
restricted. The
Congo (Zaire),
and the
species
western
closely
is
resembles
reported
by
a
have such
in the North,
and also in East Asia.
distribution, however,
Angola, Zambia,
have
Airy
P.
in the North of Thailand and Cambodia mentioned
welwitschianus Mueller
The second African
Hoffmann,
Moss)
in the Flore General de 1’Indochine
area, occurring
P. beillei. Its
growing
and he transferred the
described from African material.
(P.
specimens occurring
Phyllanthus
Hasskarl
by
peculiar pistil
consequently
feuilles plus grandes). It is remarkable, although
disjunct
described
genus beside the genus Phyllanthus.
from
separately
Shaw
P.fasciculatus
beillei Hutchinson
Hoffmann and P.
specific
Airy
name
name
P.fasciculatus
taxon
described
shortly
from the West of Java,
with other
with
to
the
Shaw
monotypic
Phyllanthus gracilis
reported
Phyllanthus
This
of
of
Hong Kong.
to
Unfortunately Roxburgh (1832)
genus.
correct
as
from East Asia
reported
name
Sprengel (1826) recognized
and
new
new name
considered the features of the
maintain the genus Ceramanthus
to
rather
was
as
genus under the
Afterwards
Phyllanthus.
and he introduced the
group of Phyllanthus
special
monotypic
anymore available because of the total different
not
Poiret
differentfrom
Phyllanthus
described in this
species
introduced it
(1790)
fasciculatus,
taxon
Sprengel
the first
was
from
part of Tanzania.
a
216
W. PUNT
Comment on the P.
Beillei-P. welwitschianus
complex
Airy Shaw and A. R. Smith (personal communication, 1971) suggest
P.
beillei and P. welwitschianus
material and believe that it all
however, there
P.
cordate base;
dried
P.
grounds. They
variable
entity.
seen
combination of
a
much African
In the material I have
seen,
elongated;
not
lateral
eight
with
(4)
nerves;
from
leaves
woody root-stocks; (2)
rounded
to
slightly
leaves often situated face
to
face in the
distinctly
a
specimens.
beillei:
cuneate
than
ovate-elliptic,
about
(3)
to one
(1) plants suffruticent, arising
welwitschianus:
oblong-elliptic
belongs
have
several taxonomic differences.
are
to
taxonomic
on
small shrubs
(1)
ft.
(3—7
towards the base and rounded
lateral
eight
(4)
nerves;
leaves
not
leaves
high); (2)
only
at
the base
elongately oblong-elliptic; slightly
(not cordate); (3) usually
situated face
strikingly
to
more
face in the dried
specimens.
On the other hand A. R. Smith wondered whether these differences make the
taxa
really good species
fires
maintaining
leaves)
in
become
some
a
or
whether it is
simply
the sucker-shoot stage
areas, whereas in other
shrub
fully-developed
or
a
question
(“typical” welwitschianus,
not
areas
small
tree
subject
two
of the biotic factor of savannah
to
with the cordate
such fires the
with rounded-cuneate
is able
plant
leaves
crown
to
(“typical’
beillei).
It is true, that both
stances.
from the
reported
Eastwards the
the
taxa
apparently
The distribution of both
eastern
lowlands
is found in
Kenya
part of Africa the species
as
well
as
at
higher
south of the equator, viz.
under
quite
species is, however,
not
states; e.g., Sierre Leone,
west-coast
species
occur
the
south
occurs
elevations. P.
Angola,
to
up
plant
quite
to
not
same.
circum-
P. beillei is
Ivory Coast, Cameroon,
Zanzibar
Rhodesia. The
welwitschianus is
is
ecological
the
eastern coast at
southern part of
southwestern part of Tanzania. The
identical
plants
reported only
etc.
(Tanzania).
are
found
from the
In
in
states
Zambia and the
Congo (Zaire),
found in the lowlands, but
at
higher
altitudes.
A. R. Smith, rather uncertain about the
only
that
and with
seems
more
fairly
constant.
delicate venation than is
intermediates exist.
Owing
entities, and taking into
them
as
to
the
account
having subspecific
species
Apparently
to
be found in P.
geographical tie-up
their great
status.
level of both taxa,
the leaves of P. beillei
(See
also
on
similarity,
Appendix
can
find
are more
one
welwitschianus, but here again
the distribution of the
two
A. R. Smith would favour
1:
feature
membranous
Note
by
A. R.
regarding
Smith.)
Sections
The four
Cluytiopsis
species
have been
Mueller
Arg. (syn.
thus Bâillon; P.
placed
in three different sections; viz. P.
Cathetus
(Loureiro)
Mueller
Arg.); P.
cochinchinensis in
albodiscus in Ceraman-
welwitschianus and P. beillei in Anisolobium Mueller Arg.
Bentham and Hooker
(1880)
maintain the three sections, but
they point already
to
the
MORPHOLOGY AND TAXONOMY
POLLEN
close
Pax and Hoffmann
relationship.
place
OF CERAMANTHUS
(1931), although
not
the sections Anisolobium and Ceramanthus just after
Cathetus
Airy
(i.e.
section
Shaw
(1969)
and
Cluytiopsis
was
strongly
BAILLON S.L.
mentioning
one
the
affinity
relationship,
any
another and the section
Paraphyllanthus together)
attent to
217
is
placed nearby.
of P. albodiscus and P. cochin-
welwitschianus. He retained, however, the sectional
chinensis with P. beillei and P.
boundaries between Ceramanthus and Anisolobium.
the
Comparing
descriptions
number of characteristics in
(1)
of the three sections, it is clear that
Some of the
common.
large (about
Flowers rather
2
mm)
in
they
important
most
have
quite
a
ones are:
with the flowers of other
comparison
Phyllanthus species.
(2) Sepals
in both
Disk in both
(3)
which is not at all
6.
sexes
sexes
present and in all sections the female disk is urceolate,
common
(4)
Stamens 3.
(5)
Filaments
(6)
Anthers free.
(7)
Anthers
(8)
All
joined
in the genus Phyllanthus
into
dehiscing longitudinally.
plants
shrubs
are
or
under-shrubs. The
disk consists of six separate segments;
three sections;
(c)
whorl differ in
shape
of the
One
same
may
most
to
or
are
shape
separate
opinion
and also the
which
below
the
shapes
might
of the
styles
are
different in all
of the
sepals
Cluytiopsis
one
important enough
styles
of the
sepals
be of value
outer
the
sepals
seems
to
in the four
seems
less
to
make three
species
are
certainly
important.
be the difference in either
in the male flowers. However, the similarities in other
and sometimes
to
are
of the
uneveness
disk-segments
so numerous
unite the three sections
The
the
(a)
and size.
only character
features
shape
are:
in the other sections the male
from those of the inner whorl. In the section
wonder ifthe differences mentioned
of minor importance
urceolate
urceolate,
form and
(b)
differences
important
in the sections Anisolobium and Ceramanthus the
different sections. In
my
The
(Webster, 1956).
column.
a
disk of male flowers in the section Ceramanthus is
are
feature
a
so
striking
that I advise,
on
taxonomic
grounds,
section.
palynologjcal results, together
with the nomenclatural consequences, are discussed
(p. 226).
DESCRIPTION OF THE POLLEN GRAINS
Phyllanthus
cochinchinensis type
Pollen Class: Pollen
arranged
in areoles
Apertures:
most common
resembling
(Table 1)
grains pantoporate
(Webster
or
pantocolporate;
sometimes the apertures
1956).
The number of
apertures
number. Ectoapertures
-
varies from 8
to
23
(rarely
up
to ca.
30)
with
usually irregular, slightly elongated pori,
the lumina of the reticulum; these apertures sometimes small, rather
in
12
as
shape
narrow
or
218
W.
TABLE
PUNT
I
Differentiatingcharacteristics
Name
Ectoaperture
1
1
Thickness of
Columellae/
the
the
capita
exine
of
of
Fusing
2
Size
3
3
the muri
00
(M)
P. albodiscus
P.
P
ca.
2.5
2.S
=
1(30-37
1 (30-37
dist.
dist.
m)
P.
P.
P-C
P-C
fasciculatus
fasciculatus
ca.
2
2
longer
at
at
top
(21-
sm
26
P.
P.
“
”
stolzianus
"stolzianus”
P-C
P-C
ca.
3
dist.
longer
longer
m)
sm-1
(22-38
(22-38 ß)
m)
P.
beillei
beillei
C
s.s.
ca.
2
utmost
utmost
=
=
top
sm
sm
28
P.
welwitschianus
C
ca.
ca.
2.5
dist.
dist.
=
=
1 (26—
1(2647
1
=
P
=
C
porus-like;
(21—
(21m)
m)
colpus-like.
2
=
about
as
long as.
3
1
=
large; sm
same
showing
are
small.
colpi (P. welwitschianus). Margin
narrow
of the
=
any
composition
special
as
features.
Endoapertures
rather broad and which have diffuse
the ectoapertures, but
of the apertures formed
the muri of the reticulum.
slightly larger
small,
-
margins.
Pori
Apertures
slightly elongated
Ornamentation: Pollen
coarse
(lumina
ca.
1.5
ß)
fused in the upper part,
Lumina
in
usually
smaller than the
always
with distinct
(lumina
ca.
2 ß
or
long
as
as
fused into
(lumina
more).
Muri
a
the
in
not
costae
length
which
of
capita. Capita
reticulum.
ca.
1
ß)
shape, always
wider than the muri
rather
to
thin, simplicolumellate,
columellate in the lower part; thickness of fused part
irregular, angular
mums
thicker than the nexine. Columellae
reticulate. Reticulum fine
coarse
sunken and
not
pori
optical section, laterally
grains
or
continuous
circular
distinct, usually longer than their capita, sometimes about
to
a
than the width of the latter.
Exine: The exine varies in thickness. Sexine
circular
by
breadth;
varying.
width of the
lumina varying.
Shape: Shape
optical
on
the
of the
pollen grains irregularly cylindrical
section the outline differs from
position
to over
from 2
to
3.5
irregularly spherical.
irregularly
circular
In
depending
in the slides.
Measurements: Longest axis
from 8
irregularly elliptic
or
to
30.
ß.
Endoporus
varying
from 20
diameter
Width of lumina
varying
varying
to
40 ß. Number of apertures
from 2
from about 1
to
ß
3 ß. Exine thickness
up
to
3.5
ß.
varying
varying
POLLEN
MORPHOLOGY AND
TAXONOMY OF CERAMANTHUS
BAILLON S.L.
219
Species
P. albodiscus
Teysman
(Ridley) Airy
Anno
s.n.,
herbarium
Java,
Java
1860,
U
s.n.; s.c.,
Number of apertures
Shaw
long
in
as
or
P. cochinchinensis
distinctly longer
P.
p.
than
Chase
To
varying
Rhodesia
(BR);
in
ca.
in
2 p, up
to
optical
329,
Cote
3.5
colpi.
as
as
d’Ivoire
capita,
Ornamentation nearly
the
grains
are
more
muri
distinctly
Columellae about
p.
section.
Longest
as
axis 30—37
p.
1—3)
usually
1
ca
China
(L.).
of ectoapertures
p,
sometimes up
spherical
p.
of P.
to
or
to coarse;
Zambia
3688,
(Cotype
in
optical
irregular, porus-like
rather
to
to
1.5
p.
section.
coarse;
muri
Columellae
Longest
axis
but
20,
very
muri
(BR);
—
or
of
Exine 2.5-3.£
colpi.
p
fused in the upper part; lumina
capita; capita small, spherical
Distinctly irregular
to
735,
27.
Cote
ca.
of the
d’Ivoire
Ectoapertures
to
slightly
in
shape.
to
rather broad,
coarse;
(WAG)
narrow
muri fused in the uppermost
1.5 p, sometimes up
slightly longer. Capita
spherical.
(Type
(Plate III, 3—7)
from 12
pilate. Shape
1986
in the range of 11—14.
rather broad
distinctly
axis 22—38 p.
J. J. F. E. de Wilde
varying
usually
short,
Columellae longer than
Longest
Graham
(BR);
stolzianus) (WAG).
Exine about 2 p thick. Reticulum rather
long
irregular, porus-like,
coarse;
2.5
to
optical
Shape
12.
part only, nearly fractured; lumina usually
about
in
rather
13658, Kwantung,
to
from 11
(beillei s.s.)
Number of apertures
short
to
shape.
1895
coarse
section.
P. beillei Hutchinson
Tehe
p,
Milne-Redhead
Stolz
apertures varying
thick. Reticulum rather
usually
of ectoapertures
sometimes up
small and
Ectoapertures irregular elongated pori,
elongate
Utrecht
(“stolzianus”)—(Plate IV)
grahamii), Kenya (L);
Number of
Shape
Plate II,
from 9
lumina
only;
Distinctly irregular
6799,
1
Kang Ping
capita; capita
beillei Hutchinson
P.
13.
to
rather broad. Exine about 2 p thick. Reticulum fine
fused in the upper part
21—26
ca
Sprengel (Plate I, 8;
1218, Hong Kong (L);
colpus-like,
from
shape.
in
Number of apertures
or
1867, Java (L); specimen
thick. Reticulum fine
p
slightly longer; capita spherical
Distinctly irregular
Coert
from 8
varying
upper part; lumina usually
capita
s.n., Anno
(L);Teysman
36876
rather broad. Exine about 2.5
fused
(Plate I, 1—7)
pollen
small and
is less
to
2
p.
Columellae
spherical. Longest
irregular
axis 21-27
than the other
species;
p.
220
W.
PUNT
PLATE I
albodiscus
Phyllanthus
section
of
(Ridley) Airy Shaw, Teysman
1.
Optical
2.
Reticulum, mud distinctly fused,
3.
Reticulum,
4.
Optical
5.
Ectoaperture, irregular, porus-like.
mud
6.
Endoaperture
7.
Section
a
more or
under the
showing
Section
showing
PLATE II (p.
irregular in
about as
as
the
columellae
which
are
higher
spherical capita.
small
and
spherical capita.
222)
Sprengel. (X
1900.)
Reticulum,
3.
Reticulum
muri
with
simplicolumellate.
ectoaperture, irregular
welwitschianus
section
5.
Ectocolpus,
6.
Ectocolpus
7.. Five
1900.)
shape.
than the
2.
Optical
(X
of 5.
high
Optical section, pollen grain irregular in shape.
4.
1860.
spherical pollen grain.
less
1.
Phyllanthus
s.n.,
cylindrical.
less
Sprengel.
cochinchinensis
Phyllanthus
or
lumina
ectoaperture
columellae
Phyllanthus cochinchinensis
8.
is more
simplicolumellate.
of
section
which
grain
of
a
narrow
in
lower
Mueller
more or
and
Drummond et al.
9.
Section
porus-like.
cylindrical pollen grain (Schmitz 284).
less
elongated.
position, distinctly
elongated ectocolpi arranged
8.
and
Arg.
in
an
fused muri
areole
7215, scanning photograph
showing thick,
short columellae
of the reticulum.
(Richards
of a
which
1077).
with
pollen grain
are
about
as
high
as,
narrow
or
colpi. (X 2000.)
slightly
shorter than the
spherical capita.
PLATE III
(p. 223)
welwitschianus
Phyllanthus
1.
Milne-Redhead
arranged
2.
in
an
areole.
beillei
3.
De Wilde
4.
Reticulum,
Arg.
of the
Hutchinson
735, reticulum,
muri
of a
pollen grain showing short,
rather broad
ectocolpus
s.s.
muri
(X
in
higher magnification. (X 7500.)
1900.)
fused at the
only
top.
simplicolumellate.
5.
Tehe
6.
Ectoaperture, colpus
7.
De Wilde
329, ectoaperture, colpus-like,
short
and rather broad.
position.
at lower
735, optical section,
exine thin with low columellae
and small
capita, shape
grain rather regular.
PLATE IV
(p. 224)
Phyllanthus
1.
Graham
beillei
Hutchinson
1986, optical
(“stolzianus”). (X
section
1900.)
irregular pollen grain.
Chase
3.
Ectoaperture, colpus-like, irregular
4.
Section
5.
Milne-Redhead
6.
Stolz
7.
Reticulum,
muri
simplicolumellate.
8.
Reticulum,
muri
distinctly fused, irregular colpus.
9.
Ectoaperture, colpus-like
ectoaperture
showing high
and
of an
2.
6799,
colpi
(X 2000.)
Scanning photograph
Phyllanthus
Mueller
8848, scanning photograph
reticulum,
columellae
3688, optical
1895, endoporus,
and small
section
of
a
circular and with
and
muri
distinctly
fused.
and rather broad.
irregular.
spherical
rather
to
slightly elongated capita.
large irregular pollen grain.
distinct costae.
of the
pollen
POLLEN
MORPHOLOGY AND
PLATE I
TAXONOMY OF CERAMANTHUS
BAILLON S.L.
221\I
W. PUNT
222\II
PLATE II
(For legend
see
p.
220.)
POLLEN
MORPHOLOGY AND
PLATE III
(For legend see
p.
220.)
TAXONOMY OF
CERAMANTHUS
BAILLON S.L,
223\III
224\IV
W. PUNT
PLATE IV
(For legend
see
p.
220.)
POLLEN
MORPHOLOGY AND TAXONOMY
P. welwitschianus Mueller
Baum
648, Angola (BR);
Zambia
7215,
Zambia
and P.
(BR);
Schmitz
Taylor 8848,
R. G.
M.
Tanzania
from 13
varying
in areoles. Exine 2—3
usually arranged
POLLEN
in
in
Distinctly irregular
Pollen
of the four
grains
exine characters
are
ectoapertures which
Webster
(1956)
are
short,
be
in P.
and Erdtman
show
compared
the New World
than 30, but
which
can
be
section
to
2
coarse
species
a
are
can
(1952).
only
a
in minor
in the
from
points
capita
usually
species
to
species.
distinctive differences
most
but distinct
colpi
and
found in the
are
less
more or
regularly arranged
be
show any
not
compared
Areolate
regular arrangement.
with the areoles
pollen grains
are
pollen grains
of the areoles
colpus
have their
regular
by
known from many New
sandwicensis, P.
as
the
virgatus,
welwitschianus
pollen grains
in this
like in P. welwitschianus.' In
endoapertures
the ends of
at
(Webster, 1956).
fused in the uppermost part
are
The
described
the ornamentation in P.
opinion
in the middle of the
angles
as
the columellae are
only,
(1)
in
while in P. albodiscus
distinctly larger
than the capita in
as
long
as
this difference is,
the
capita;
however,
not
importance.
fasciculatus
resembles in many respects P. beillei and the
a
rather
beillei shows
to
“
as
n,
and the
great similarity in shape and size,
and from the section Macraea (P.
distinctly fused; (2)
P. beillei shows
P.
long
axis 26—47
great similarity in shape, apertural characters
which do
the areolate
found in the reticulum. P.
like
;muri distinctly
Columellae as
ju.
spherical. Longest
cochinchinensis is smaller in size than P. albodiscus,
of much
P.
slightly broadened,
or
P. cochinchinensis, while in P. albodiscus the columellae are about
P.
1077,
Milne-Redhead
in the range of
usually
narrow
cochinchinensis differs from P. albodiscus in several characteristics:
the muri
P.
Richards
welwitschianus, whereas the ectoapertures in P. albodiscus
endoapertures
P. cochinchinensis the muri
(3)
and Rutherford-Smith
(BR);
547, Congo (Zaire) (BR);
best with the areoles in the section Macraea,
syncolpate colpi
P.
Zambia
thick. Reticulum rather
differ
in the Old World. In my
section have their
the
Drummond
278,
sometimes up
optical
narrow
species (Webster, 1956)
etc.) occurring
can
/u
ju,
pollen grains
arrangement in P. welwitschianus
World
Detilleux
to more
very much alike. The
6-angular figures
or
Richards
shape.
species
irregular, poms-like apertures
are
(BR);
welwitschianus and P. albodiscus show
in 5-
225
MORPHOLOGY
and ornamentation. The
P.
1.5
ca
slightly longer. Capita robust,
ju.
S.L.
(BR).
fused in the upper part; lumina
about 35
Zambia
Ectoapertures always colpi
16—20 apertures.
or
Young 1047, Angola (BR);
13/7,
284, Congo (Zaire) (BR);
Number of apertures
BAILLON
Arg. (Plate II, 4—9); plate III, 1—2)
Mitchell
(BR);
OF CERAMANTHUS
cochinchinensis has
coarse
pollen grains
call for convenience P.
stolzianus
to coarse
of
two
a
fine
to
rather
most
coarse
striking
difference is
reticulum,
while
reticulum.
different kinds. An
“stolzianus” and
a western
eastern
one
kind, which I should
which I call P. beillei
s.s.
”
is
easily recognizable
by its thick exine, distinctly fused muri, irregular,
W. PUNT
226
very short, broad
with
grains
which the muri
the
shape
is
elongated pori
less thick exine,
a
species
”
This reticulum is
resembles P.
help
hand,
P.
do
show
not
nearly retipilate;
common
so
pollen
reticulum of
a
fasciculatus
in
quite
a
slightly
with the
pollen
much resemblance
to
the
solving
unlike the
s.l.
According
more
or
to
specimens, however, certainly
of less
is
features,
On the
shows
specimen
not one
importance
may be of
supposition, pollen morphology
“welwitschianus” and “stolzianus”. Also the
less intermediate
pollen grains (Richards 278,
this
in the taxonomic
pollen grains intermediate between
problematical,
All
Apparently pollen morphology
in the P. beillei-P. welwitschianus complex.
problems
tendency
of P. beillei show differences
specimens
western
specimens (P. “stolzianus”).
beillei.
the
of the
pollen grains
eastern
species
in
has
s.s.
the much thicker exine and the
by
pollen grains
but its
taxon, viz. P. beillei,
to one
within the
P. beillei
in the minor details.
with those of the
other
only.
stolzianus
but differs from it
species,
It is remarkable that
little
“
P.
only slightly irregular.
of the other
belong
distinctly irregular shape.
a
reticulum. P. beillei s.s. has the main characteristics in
coarser
other
and
short, rather broad, but distinct colpus and
fused in the uppermost part
are
number of characteristics,
grains
a
show either
specimens,
1102)
Robson
or
typical
typical “welwitschianus”
“stolzianus”
pollen grains (Faulkner
3165).
small differences
Although
is
it
species,
obvious that
type. However,
the
same or
and
not
one
they
all
between the
belong
to one
same
any of them show
seen,
characters. I have
seen
of the
pollen grains
the P.
of the four mentioned
pollen grains
entity,
wonder if there is
other
may
any
almost the
material I have
occur
the
species
over
same
Phyllanthus
300
in this
species
kind. As far
as
cochinchinensis
large
I
genus showing
to
up
can
cochinchinensis type stands rather
and
apart
this
judge
cannot
moment
from the
be
with any other
the genusPhyllanthus.
type in
compared
CONCLUSIONS
(1)
The taxonomy of the four
maintain three sections. The
species
does
not
only
value is present in the male flowers of P. albodiscus
disk is
more
perhaps
strong argument
characteristics in
of
consisting
(2)
the
a
The
same
(3)
to
to
differentiating
one
uphold
There
species
are
two
(sect. Ceramanthus).
distinct sections,
sections,
which
laid
(1956)
and
as
he
of the
most
so
much
The urceolate
there
although
always
have
a
are
many
male disk
more
that
they undoubtedly belong
on
the
characters
to
pollen morphology
pollen
and sections it
characters
seems
to
sufficiently
are not
types. These minor differences, however,
mentionedthe
subgenera
important
alike,
can
species.
great emphasis
usually
characters for
are
minor differences which
only
differentiate them into
Since Webster
as
of all
differentiate between the
genus Phyllanthus
results
to
disk-segments.
pollen grains
pollen type.
important
be used
to
with the other
common
fundamental differences
give enough
differentialcharacteristic which may be of sectional
as
justified
within the
one
of the
to use
unite the three sections.
the
pollen
POLLEN
MORPHOLOGY AND TAXONOMY
and
(4) Taxonomy
leads
pollen morphology
Ceramanthus and
Anisolobium,
OF CERAMANTHUS
nomenclatural reasons this section has
to
me to
have
Cluytiopsis
to
BAILLON S.L.
227
the conclusion that the sections
be united in
one
section. For
large
be named Ceramanthus Baillon.
ACKNOWLEDGEMENTS
The author wishes
he is
particular
express his sincere thanks
to
to
Dr A. R. Smith
for his valuable advice and for his kindness
Gardens, Kew)
grateful
Professor Dr F. P. Jonker for
to
send
to
the
criticizing
Elsendoorn for his
in
help
photographical
typewriting. Finally
Kew and
Leiden,
he is much indebted
Wageningen
matters;
Mrs Ch. M.
to
to
van
to
Mr T.
Schipper
Wessem-Alvares
He is
Mr H. E.
to
for his skillful
for her careful
Vega
the directors of the herbaria
for their kindness in
material. In
manuscript.
indebted to Mrs G. Coomans-van der Mark for her technical assistance;
arrangement of the plates and
Botanic
(Royal
interesting
at
Brussels,
the author with all the
supplying
material he asked for.
REFERENCES
Airy Shaw,
23:
H.
1969. New
K.,
noteworthy species
or
of
Phyllanthus
L. Kew.
Bull.
(R.
Bot.
Gardens),
26-40.
1858. Etude
Bâillon, H.,
générale
du
Groupe des Euphorbiacées.
Libraire
de Victor
Masson, Paris,
684 pp.
Beille, L., 1927. Phyllanthus.
Paris,
5:
Bentham,
De
G. and
Loureiro, J.,
Erdtman, G.,
J.
Hooker,
1790.
P.
C,
Pax,
1844.
Palynol.,
F. and
J.
748
L.
M.,
Genera
Cochinchinensis.
Morphology and
Catalogus
Flore
générale de Undo-Chine. Masson,
Plantarum, 3(1).
Ulyssipone,
Plant
745
Heeve
and
459 pp.
Co., London,
pp.
1.
Taxonomy.
Angiosperms. Almqvist
and
Wiksell,
Plantarum
in
Horto Botanico
Bogoriense
Culturam Alter.
Lands
391 pp.
Methode
Herstellung
zur
von
Schnitten
durch
Pollenkörner.
azetolysierte
8: 252-254.
1931.
Hoffmann, K.,
Natürlichen
Poiret,
1880.
D.,
1968. Eine einfache
Grana
(Editoi).
539 pp.
Drukkerij, Batavia,
Leins,
J.
Flora
1952. Pollen
Stockholm,
Hasskarl,
In: M. H. Lecomte
571-608.
Euphorbiaceae,Phyllanthus.
Pfanzenfamilien. Engelmann, Leipzig,
1804. In:
J. Lamarck
2nd
In:
ed,,
A.
Engler
pp.
and Harms
(Editors),
Die
60-66.
(Editor), Encyclopedic Methodique Botanique,
5.
Agasse, Paris,
pp.
Praglowski, J.,
grains.
1970.
The effects of
Rev. Palaeobot.
1967. Pollen
Punt, W.,
Palynol.,
Reitsma, Tj.,
3:
grains.
morphology of
1969. Size
1970.
Rev.
Roxburgh, W„
Sprengel, K„
Webster,
G.
modification
9:
and the
embedding
nedia
on
the
shape
of
pollen
203-208.
the
genui
Phyllanthus (Euphorbiaceae). Rev.
Palaeobot.
Suggestions
1832. Flora
of recent
towards
Indica,
10:
unification
3. Allen
monographie study
Arbor. (Harvard Univ.),
under
different treatments.
Rev.
37:
of
descriptive terminology
of
Angiosperm pollen
39-60.
Systema Vegetabilium,
1956. A
pollen grains
175-202.
Palaeobot. Palynol.,
1826.
L.,
10:
141-150.
Palaeobot. Palynol.,
Reitsma, Tj.,
pre-treatment
Palynol.,
217-256.
and
Co., London,
3. Librariae
655 pp.
Dieterichianae, Göttingen,
of the West Indian
species
832 pp.
of Phyllanthus. J. Arnold
228
W.
APPENDIX
Note
by
A.
1
R.
on
of Punt’s
216,
p.
Smith'
extreme forms
Although
matter
and which
it is not
the
for
example,
unbranched, arising
face),
has
which
some
more
than
appear
with
a
of P.
a
the 9
out
for P.
species
we have
the
leaves
pollen,
of lateral
shoot of Robson
1102,
278 from northern
beillei,
beillei; placed
and
represented
weigh
he
the leaves
side
by
side with
without
a
is
the low
somewhat
stature
and
a
is
a
d
Faulkner
doubt,
(30
botanist
3165,
Botanic
base;
or
cm
of
shoots
leaves
height)
species.
only
reliable
guide
states,
species they
Malawi,
situated
with
in
which
less
face to
the 6
at the
shoot,
base;
and the fact that it
There
does thus
might be forgiven if, presented
we
again
eight pairs
at the
from
have
of lateral
base,
a
mixture
nerves
which would
Zanzibar, in
difficult at first
which
sight
to
and
to the
separation of
Gardens, Kew, Richmond, Surrey (Great Britain).
P.
of features.
are
the leafbase
appreciate
beillei
not
tend to rule it
here.
is the
indicated
more or
less rounded
in
subject
beillei rather than P. welwitschianus.
cordate
it is very
to which
(i.e.
this latter
specimen,
have more than
are
at the
he has
intermediate
Manning District
or more
against
it with P.
which
represent
of the
as
undoubtedly conspecific
leaves cuneate
placed
which
P. welwitschianus.
Royal
form part
grounds alone,
cordate
which
shoot,
face; nevertheless, they
Perhaps pollen morphology
1
which
of P. welwitschianus
dimorphism,
Zambia,
these
perceptibly
the 9
nerves), although
tend to
on
1102 from the Fort
(i.e. bushy habit;
and which is P. beillei
are
Robson
rootstock;
by
certainty,
characteristic features
beillei
situated face to
Phyllanthus species
problematical specimens
with
certain amount of sexual
The habit is that of P.
rounded,
a
more
to say
woody rootstock,
In Richards
strikingly
exist
the
out
eight pairs
to be
only
has
from
is borne
features
arises from
there
possible
so
d shoot
of the two African
fairly readily distinguishableon morphological grounds
are
paper
nevertheless
belong. Thus,
although
PUNT
and
is
that two