A JOURNAL ON TAXONOMIC BOTANY,
PLANT -SOCIOLOGY AND ECOLOGY

REINWARDTIA

. Editors
MIEN A.

KIJS WAT A KARTAWINATA
N. WULIJARNI-SOETJIPTO

1 Published by '
HERBARIUM BOGORIENSE

LEMBAGA BIO-LOGI NASIONAL — LIPI
BO.SOR, INDONESIA

Eeinwardtia Vol. 9, Part 1, 1 —182 31 December 1974

10- ISSN 0(f34-365X



1! El N WARD TI A
Published by Herbarium Bogoriense — LBK, Bopor

Vol. 9, Part 1, pp. 153 —I76 (1371)

POLLEN MORPHOLOGY OF CERTAIN TROPICAL PLANTS

A. N. RAO & LEONC FONG LING
Department of Botany, University of Singapore, Singapore

ABSTRACT

The pollen morphology of 49 species belonging to 48 genera and 31
families ia recorded in this paper. Of these 24 anil 19 taxa are studied

show the variations in the local pollen grains. In most of the species
studied presently the characters were similar to those recorded previously
for the respective families. The variations seen in some of the taxa with

The present observations ace comparatively discussed with reference to
previous literature.

ABSTRAK

tei-Morfologi serbuk sari 49 jenis tumbuh-tumbuhan tropika
golong dalam 46 marga dan 31 suku telah dilaporkan dalam karangan ini.
Enam jenis telah dipertelaken kembali untuk menunjukkan adanya variasi
pada serbuk sari tumbuh-tumbuhan setempat. Ciri-ciri kebanyakan jenis
yang diteliti ternyata sama dengan ciri-ciri sukunya yang bersangkutan
yang sudah dilaporkan orang aebelumnya. Variasi yang terlihat pada
sifat-sifat dinding serbuk sari, keadaan lubang-Iubang dan Iain-lain di-
tunjukkan. Hasil pengamatan yang dilakukan sekarang ini dibandiugkau
dan dibahas dengan mengafu pada puhtaka-pustaka yang ada.

IXTRODUCTtON

According to a recent estimate there ftre about 28,000 species of
angiosperm in the Malesian region representing about one tenth of the
world flora (Van Steenis 1948). The estimated number of Malayan
genera is around 1,800 and of Malayan species about 8,300 (Keng 1970).
Taxonomieally these are well described and documented in the different
regional floras (see Keng 1970, for details). For the last few years
detailed investigations are tarried out to study the pollen morphology
and other structural details of some of the species. In an earlier publi-
cation the pollen morphology of about ninety local species, representing
forty families of angiosperms, was described (Rao & Lee 1970). The
Pollen characters of about 50 species are recorded in this paper, and few
of the tasa are described to indicate the variations seen in the local
plants.

— 158— . . . .



REINWARDTIA [VOL. 9

The well known works of Cramvell (1953), Erdtman {1943, 1952,
1957, 1961), Faegri & Iveraen (1950, 1964), Hyde & Adams (1958), Nair
(1965) and Wodehousc (1935) and the journals Grana PalynologiRil
Botanical Review (Faegri 1956; Wodehouse 1936), Pollen et Spores^
Palaeobotanist and Journal of Palynology served as chief sources of
references. Other published papers were consulted wherever necessary.

2O/J

^^K .1974]

Pigs. 1-8. 1. Imperata cylindrica. 2. Schizostachyum. braekycladum. S. Arckonto
phoenix alexandrite. 4. Aglaonema jrictus. 6. Aechmea tinetoria. 6. Pitcairn.it

iategrifoUa. 7. Gnnocaryum Utorate. 8. Boerhaavia diffusa.

MATERIALS A NO METHODS

Pollen grains were collected in the mornings from the fresh ope
flowers of plants growing in Singapore and West Malaysia. Envelope
were used to separate and prevent contamination of the pollen of diffe

RAO & LEOKG: Ti-fipicat pntten grai;

rent species. In cases where the anthers were large (such as Datura)
one flower gave all the material required whereas in others (such as
Grossandra and Nyctantheg) 15—20 flowers were found to be necessary.
Where contamination of foreign pollen was found, due to the visiting
pollinators, the very mature buds were used. Voucher specimens were
prepared and deposited in the herbarium and the numbers of such speci-
mens are mentioned against the species name. The symbols USH (Uni-
versity of Singapore Herbarium) and SBGH (Singapore Botanic Gardens
Herbarium) indicate the place of deposition and the number mentioned
here is the same as recorded on the herbarium sheet.

The pollen grains were fixed in concentrated acetic acid and after
24 hours they were acetolysed following the method outlined earlier
(Erdtman 1952, 1960). The procedure was slightly modified as was
found necessary. In the case of thick walled pollen grains it was necessary
to bleach them before acetolysis, by using saturated sodium chlorate
solution.

To obtain the average measurements at least 20 grains were measured
in each case. The size of fresh and acetolysed grains was compared and
there was insignificant difference between the two. To determine the
size, shape, L0 pattern, exine pattern, the methods employed by earlier
workers were followed and descriptive terms used presently are in con-
formity with the published work (Erdtman 1952; Nair 1965). In the
palynograms of dicotyledonous plants the first figure on the left repre-
sents the polar view, followed by a second of the equatorial view, with
the exceptions of the pantoporate or rugulate conditions. For monocoty-
ledonous plants the first diagram represents the surface view followed
by a diagram showing lateral view. The OL patterns are given except
for those where sexine pattern is psilate.

OBSERVATIONS

MONOCOTYLEDONS

G K A M I N E A E

Imperata oylindrica Beauv. USH 4033. Figure 1 Plate I, 1

Monoporate, pore diameter 2.1 .̂m, pore margin 1.6 .̂m. Exine
1-1 y.m thick. Spheroidal, diameter 27.4 jim. Foveolate.

•Sekixostackyum brachycladum Kurz. USH 5279. Figure 2

Monoporate, of pore diameter 2.4 iim, pore margin is thick forming
an annular ring of 1.8 iim. Exine is 1.3 jim thick. Spheroidal, diameter
28.1 jim. Foveolate.



REINWARDTIA [VOL.
1974] RAO & LEONG: Tropical pollen, gr

In all the genera so far studied, pollen grains are monoporate and
differences are noted in the diameter of the grain, the porate condition,
that is whether they are crassimarginate or tenuimarginate.

P A L M A E

Archontophoenix alexandrae H. Wend] & Drude. SBGH 15/15. Figure 3

Monosulcate. Exine 0.9 jun thick. Bilaterally symmetrical 42.3 x
49.5 X 28.8 ij.m. Psilate.

This family ia palynologically heterogenous (Erdtman 1952). The
OL pattern ranges from psilate to reticulate, some of them beset with
spines and others not so. Generally, members of this family are either
1-sulcate or 2-sulcate.

A R A C E A E

Aglaonema pictum Kunth. USH 623. Figure 4

Non-aperturate. Exine thickness of 3.6 [AIH. Spheroidal, diameter
29.8 nm. Exine plicate, that is folded. OL pattern foveolate.

This is a palynologically heterogenoua ranging from the non-apartu-
rate to the sulcate and porate conditions. In this species, the exine is
unique because it is plicate and its OL pattern is more or less obscure.
Such non-apcrturate pollen grains with thick plicate exine is considered
more "advanced" than the monosulcate, finely reticulate ones (Erdtman
1952).

B R O M E L I A C E A E

Aeckmea tinctoria Mez. USH 4458. Figure 5

Biaperturate, one pore at each end of the pollen grain. Exine l.-l
ijm thick. Bilaterally symmetrical 22.1 x 34.9 x 24.2 y.m. Reticulate.

Pitcairnia, iutegrifolia Hort. SBGH 17/171. Figure 6 Plate I, 2.

Monosulcate. Exine thickness 1.0 nim. Bilaterally symmetrical 19.2 X
32.6 x 21.0 urn. Retipilate.

In Bromelioideae, 1-aperturatc grains are found and Aeckmea
involucrata belonging to this sub-family is not an exception (Erdtman
1952). But Ae.chmm tinctoria is exceptional in that the pollen grains
are biaperturate. Pollen grains of another species A. bernoulliana show
the biaperturate condition also (Erdtman 1952). Here, as in pollen
grains studied in certain species of Ochnagavia, Thecophyllwm, etc.

(Erdtman 1952), the ends of the grain, that is around the regions of
the apertures show a finer reticulation than the rest of the grain surface.
Members of the group Pitcairnioideae have monosulcate pollen and
pitcairnia. iutegrifolia is no exception (Erdtman 1952).

P O N T E D E K I A C E A E

Eickkornw, crassipes Solms. USH 4358. Figure 10

Monosulcate. Exine 1.5 ii.ni thick. Bilaterally symmetrical. Renate,
20.5 x 43.1 x 23.3 pn. Finely foveolate.

Monoekorut hastata Solms. SBGH 4/179. Figure 11

Monosulcate. Exine thickness 1.2 -im. Bilaterally symmetrical
17.6 x 37.4 x 18.9 jtm. Foveolate.

The pollen grains of various genera show sulcate condition.
Essentially the pollen grains of Eichhornia crassipes and that of Mono-
choria hastatv are similar except for the slight difference in size.

grandiflora. 10. Eichhornia. craaaipee. 11.
a. 13. Zephyranthes alba. 14. GdustacKys



REINWARDTIA [VOL. jj RAO & LEONG: Tropical pollen

A M A K Y I , L I D A C E A G

LeptocMlon qia'toensis Sealy. USH 4459. Figure 12 Plate I, 3
Monosulcate. Exine 2.4 ;J.ra thick. Bilaterally symmetrical 49.4 x

79.2 x 46.9 urn. Retipilate.

Zephyranth.es alba Hort. USH 4364. Figure 13
Monosulcate. Exine thickness 1.2 am. Bilaterally symmetrical

27.4 x 51.6 x 27.1 ^m. Very prominently reticulate.

Pollen morphology of members of this family are diversified
(Erdtman 1952). Some of them are small and in others very large
pollen grains are found here. Longest axis 17 to 160 ;im in some "rains
(Erdtman 1952). Likewise, Oh pattern ranges from scrobiculate to
retipilate. But generally, the aperturate condition is sulcate. Erdtman
has divided this family into four main groups on the basis of pollen
morphology (Erdtman 1952).

Z l N G I B E E A C E A E

Geostackys rupestris Ridley. SEGH 12,-170 No. 8. Figure 14 Plate I, 4
Non-aperturate. Exine 0.6 nm thick. Spheroidal. Diameter 87.2 ym.

Psilate, but excrescences are in the form of gemma.
Pollen grains of members of this family are usually non-aperturate,

large in size, but generally, the exine is very thin and hence not resistant
to aeetolysis. Exine may be psilate, and sometimes excrescences may be
present in the form of spinules or gemma.

DICOTYLEDONS
N Y C T A G I X A C E A E

Boerhaavia diffiisa L. USH L71 '71. Figure 8

Pantoporate. Pore diameter 2.4 j.m. Pore margin 1.1 ;i.m. Interposal
distance is 17.1 ;im. Exine 5.4 .̂m thick. Spheroidal, diameter 60.3 |ini.
Surface of pollen grain is spiniferous, with spine length 2.4 \j.iti. Inter-
spinal areas foveolate.

Pollen grains of different genera show spines or spinules. The
aperturate condition is very varied, ranging from colpate to pantoporate
(Erdtman 1952). Pollen of another species, Boerhaavia caribaea is
studied by Lundell (cited in Erdtman 1952) and it is found that the
porate condition and the excrescences are essentially the same, except
for the fact that the pollon diameter of this species is bigger, namely
80 jim. This is true for many other species studied by Nowicke (1970)
e.g. B. anistyphyUa, B. dichoto'nia, B. erecta, etc.

P O B T U L A C A C E A E

ica grandiflora Hook. USH 2711. Figure 9

Polyrugate. Exine thickness 2.3 ;j.m. Spheroidal. Surface of the
,rain spiniferous, with spine length 1.4 u.m. Interspinal area areolate.
fhe arrangement of the colpi is irregular and there are about five in
sch pollen grain. Oil globules are also seen in the grain and they vary
om two to four in number.

Pollen of Portidaca quadrifida and Portidaca grandiflora, a cultivated
species in Uppsala, are described (Erdtman 1952). In all cases, the
polyrugate condition is seen, but spinules of Portulxiea, grandiflora grown
in Uppsala are twice the length of that of the same species grown locally.
This difference could be attributed to environmental conditions as the
climate in Sweden is very different from that of Singapore (Kurtz &
Linerman 1958).



REINWARDTIA

CRUCIDERAE

[VOL. 9

Braasiea oleracca L. USH 4457. Figure 15

Tri-zonocolpate. Exine thickness 1.2 ;jm. Prolate spheroidal 14.9 x
33.4 tjm. Reticulate.

Other species studied are Braasiea campeal.ris (spheroidal, 35 ;im
and Braasiea Juneea (spheroidal, 21 ,um; Nair 1965). The pollen aperturate
conditions of these 2 species are also tri-zonoeolpate, but their sizes and
shapes differ from B. oleracea.

MORINGACEAE

Moringa oleifera Lam. USH 4454. Figure 16

Tri-zonocolporate. Lolongate ora 7.5 x 5.2 .̂m. Exine 1.2 ym thick.
Oblate spheroidal 32.4 x 33.3 ii.m. Foveolate.

Another species was studied, Moringa aptera, cultivated in Egypt
(Erdtman 1952). The pollen characters of both these species are similar,

C U N O N I A C E A E

Weinmannia blumei Planch. USH 3588. Figure 18

Tri-zonocolpate. Exine thickness 0.8 .̂m. Prolate spheroidal 10.2
9.3 jtm. Foveolate.

Pollen grains of members of this family are usually 2- or 3-colporate
with exine stratifications more or less obscure. Weinmannia intermedia,
grown in Mexico, has pollen grains of the colporate condition (Erdtmatt
1952). But Weinmannia blvmei shows colpate condition.

L E G U M I N O S A E

Andira mrinamensis Splitg. USH 739. Figure 17 Plate I, 5

Tri-zonocolporate. Pollen grains aspidote, that is the apertures are
borne on small circular areas called aspides, protruding as rounded domes
from the surface of the grain. Ora circular 1.3 ^m in diameter. Exiw
thickness 0.9 jra. Prolate 17.6 x 12.8 \ita. Finely foveolate.

There is considerable variation in the family with aperturate
condition ranging from colporate to porate and exine pattern ranging
from psilate to reticulate (Erdtman 1952; Nair 1965). The pollen in i hi -
case is monad in contrast to the polyac! condition in many Mimosoidcac

RAO & LEONG: Tropical pollen grah

2 O J J

Figs. 22-27. 22. Elaeodexdron quadrangulatum. 23. Impaliens griffUMi.
24. Wormia aiiffrutico&a. 35. Pierie ovalifolia. 26. Rhododendron leucobotrys.

27. Styrax benzoin.

LlNACEAE

Ixonanthes reticulata Jack. USH 402. Figure 19 Plate I, 6

Tri-zonocolporate. Ora lolongate 9.9 x 5.2 (j.m. Exine 1.9 j.m thick.
Prolate spheroidal 50.3 x 49.7 jum. Surface of pollen grain spinulous,
With spinule length of 0.9 ;jm. Interspinal areas foveolate.

This species was collected from Kedah Peak and when compared with
the pollen of Singapore plants studied (Geh 1967), it had pollen grains
of a larger size. This could be attributed to different environmental
conditions, as the species collected from Kedah Peak was found at an
altitude of 3,000 feet above sea level (Kurtz & Liverman 1958). The
ora was larger and lalongate compared to that of Singapore plants



162 REINWARDTIA [VOL. <!

which had a smaller and circular ora. Pollen grains of other species,
/. icosandra, and /. chinensia (colporate, spinulose) have similar morpho-
logy (Erdtman 1952).

E U F H O R B I A C E A E

Jatropha podagrica Hook. USH 2282. Figure 20 Plate I, 7

Non-aperturate. Exine thickness 2.9 ;j.m. Spheroidal, diameter
61.5 |im. Excrescences in the form of verruca. Areolate.

Pollen of Jatropha curcas has a diameter of 68 ;j,m (Erdtman 1952).

C E L A S T R A C E A E

Elaeodendron quadrangitlatum Reiss. USH 4359. Figure 22

Tri-zonocolporate. Ora lolongate 17.5 x 1.4 ^m. Exine thickness
1.7 yjn, thinner towards colpi margins. Subprolate 29.5 x 25.6 ;itn.
Punctate.

" . I C A C I N A C E A E

Gonoearyum Morale Sleum. USH 4354. Figure 7

Non-aperturate. Exine 2.6 jim. thick. Spheroidal, diameter 29.3 ;;.m.
Retipilate.

This species is an exception in that it is non-aperturate. Normally,
pollen grains belonging to members of this family have the colpate,
colporoidate, or porate conditions (Erdtman 1952).

B A L S A M I N A C E A E

Impatiens griff ithii Hook. f. SBGH 19/38C/1. Figure 23
Pantoporate with at least three pores. Pore diameter 4.5 ;im. Pore

margin 0.6 iim. Irregular distribution of pores and therefore the inlev-
poral distances are not constant. Exine 1.3 jim thick, spheroidal, diameter
34.1 jim. Spmulous with spinule length of 0.8 |im. Reticulate.

Members of this family have pollen grains of the colpate condition
and show variation in size (Erdtman 1952; Nair 1965; Faegri & Iversen
1949). But an exception is found in this species Impatiens griff ithii,
which shows pantoporate condition. Other species, namely Impatiens
biflora (24—34 |ira), /. noli-tangere (23—31 jim), and I. parviflora
(29—40 ;Am) have grains of the colp(old)ato condition. (Erdtman 1952;
Nair 1965).

1D74] LKONG: Tropical pollen, grains

D l L L E N I A C B A E

fflormia suffrutieosa Griff. USH 1747. Figure 24
Tri-zonocolpate. Exine thickness 1 ;im. Suboblate 18.6 x 22 inn.

Reticulate.

The eolpate condition is found in Dillenia ovata, Wormia alata,
fetracera alnifolia, etc. (Erdtman 1952). Sometimes, brevicolpate
condition is seen in the pollen grains of Wormia suffrutieosa cultivated
in Bogor (Erdtman 1952).

E R I C A C E A E

Pierix oval!folia D. Don. SBGH 14/93. Figure 25
Compound pollen grains in tetrads, of two types, namely the

tetrahedral and the tetragonal type i.e. 1 pair of pollen grains art
perpendicular to the other pair. The tetrahedral type occurs in 83%
and tetragonal type occurs in 17%. Tri-zonocolporate. Ora lalongate
2.8 x 4.2 |im. Exine 1.1 ;j.m thick. Oblate 17 X 23.8 M». Longest axis
of tetrad 33.9 ;im. Finely foveolate.

s arbor-tristis. 29. Cerbera odolla
.iterria. 31. Ronpellia grata.



184 REINWARDTIA [VOL. 9

Rhododendron leucobotrys Ridley. SBGH 46/93/8. Figure 26 Plate I, 8

Compound pollen grains in tetrahedral arrangement. Tri-zonoeol-
porate. Ora lalongate 2.5 x 10.5 ;jm. Exine thickness 2.0 ;im. Longest
axis of tetrad 51 Jim. Foveolate.

The pollen grains of this family are either united in tetrads, or
occurring singly. Aperturate condition is usually colporate with exine
obscure. The ora is lalongate and in this case, it is very prominent
forming a streak across the colpus. Size variations are seen in the grains
of this family, with a range between 25 jun to more than 50 ;im. Other
species studied are Rhododendron arboreum (58 jim), R. barbatum
(54 ]ini), R. catawbiense (62 ]im), and R, lepidotivm (53 ;jm). They show
similar pollen morphology except for the size variations (Erdtman 1952;
Nair 1965).

S T Y E A C A C E A B

Styrax benzoin Dryand. USH 4350. Figure 27

Tri-zouoeolpate. Exine thickness 3.1 y.m. Prolate 32.4 X 23.3 jim.
Foveolate.

The pollen grains of Styrax officuialis have the colporoidate condition
and are much larger in size. Styrax suberifolhtm has pollen grains of
the colporate condition (Erdtman 1952). Therefore, the aperlurate
condition is not a constant feature, even in the pollen grains of the same
genus.

0 LEACEAE

Nyetanthes arbor-tristis L. USH 333. Figure 28 Plate I, 9

Tri-zonocolpate. Exine 3.4 .̂m thick. Subprolate 46.3 x 39.1 FJ.HI.
Retipilate.

Pollen grains of local plants are similar in size, and it is subprolate
in contrast to the spheroidal condition (Erdtnian 1952). The OL pattern
of the pollen grains of the species found locally is retipilate and do not
possess any excrescence in contrast to that found in Thailand, which
have a reticiiiate pattern with the lumina studded with granules or
± piloid excrescences. This could be due to the different climatic
conditions which existed in different habitats or they are obtained from
different horticultural varieties (Kurtz & Liverman 1958).

1974 RAO & LKONG: Tropical pollen grains

A F O C Y N A C B A E

Cerbera odollam Gaertn. USH 384. Figure 29

Tri-zonocolporate. Ora lalongate 15 x 6.6 jmi. Exine thickness
1.8 î m. Shape ranges from subprolate to prolate spheroidal 72.8—75.3 x
61.6—72.8 ji. Foveolate.

Ramvolfia vomttoria Afzel. USH 4460. Figure 30 Plate II, 1
Tricolporate. Ora lolongate 5.6 x 3.4 ;ijn. Margin of colpus, crassi-

marginate. Exine 1.2 ;j.m thick. Suboblate 29.9 x 37.8 jtm. Foveolate.

RoupeUia grata Wall & Hook. USH 352. Figure 31
4-zonoporate. Pore diameter 2.S [im. Pore margin 1.5 .̂m. Interporal

distance 22 ym. Exine thickness 1.2 ii.m. Spheroidal, diameter 40.4 |xm.
Foveolate.

Thevetia peruviatia Merrill. USH 4355. Figure 32
Tri-zonocolporate. Ora lalongate 5.5 X 6.3 ;j.m. Exine thickness

2.8 ;j.m. Suboblate 50.5 x 62.9 iim. Reticulate.

2On

p 'gs. 32 - 36. 32. Thevetia pemviana. 33. Ekretia mv
sjieciosisffimMm. 35. Bmnfolsia etlyeiaa. 36. Br

•ophylla. 34. Clerodendr-



REINWARDTIA [VOL. 9

Pollen grains of members of this family are usually colporate or
porate in nature. Pollen grains of Rauwolfia verticillata are colporate,
and ora are lalongate compared to the lolongate condition found in that
of R. vomitoria (Erdtman 1952). The exine pattern is usually foveolate.
The main morphological characters are fairly constant within individual
genus (Erdtman 1952).

BOKAGINACEAE

Ehretia mierophylla Hort. USH 4361. Figure 31 Plate II, 8
Tri-zonocolporate. Ora lalongate 3.8 x 1.9 [im. Exine thickness

1.2 jim. Prolate spheroidal 27.1 x 27.8 y.m. Faintly reticulate.

Pollen of other species studied are Ehretia (aevis var. floribunda,
E. latifolia, E. rigida and E. silvatica (Erdtman 1952). The morphological
characters in all these species are similar.

V E R B E N A C E A E

Clerodendron speciosissimum Van Geert. "USH 4537. Figure 40 Plate II, 3
Tri-zonoeolpate. Exine thickness 2.5 yjn. Prolate spheroidal 50.6 x

46.3 y.m. Exine surface is spiniferous with spine length of 1.3 y.m.
Foveolate.

Two other species have been studied, that of Clerodendron infor~
tunatum: spheroidal, 60 urn and C. serratum: spheroidal, 71 ji.m (Nair
1965) and it has been found that the general pollen morphology conforms
with that of Clerodendron speciosissimum.

SOLANACBAE

Brunfelsla calycino, Benth. SBGH 61/114. Figure 35 Plate II, 4
4-zonoporate. Pore diameter 9.6 |im. Pore mai-gin 0.9 |im. Interporal

distance 14.6 iim. Exine thickness 1.0 ;im. Oblate spheroidal 35.8 x 35.9
um. Reticulate.

Brunfelsia wndwlata Swartz. USH 585. Figure 36
Tri-aonoporate. Pore diameter 8.4 -im. Pore margin 1.3 |im. Inter-

poral distance 23.5 y.m. Exine 1 |im thick. Oblate spheroidal 37.9 x 39-3
iim. Foveolate.

Capsicum annuum L. SBGH 10/114. Figure 39
Tri-zonocolporate. Ora lolongate 3.4 x 5.8 um, forming a long streak

across colpus. Exine thickness 1.2 jim. Prolate spheroidal 23.6 x 21.4 um.
Foveolate.

RAO & L E O N G : Tropical polleti yra 167

Datura metel L. SUH 445. Figure 37
Tri-zonobrevi col pate. Exine thickness 1.1 ji.m. Oblate spheroidal,

40.4 x 33.1 um. Striate. Striations converging at the polos.

Petunia axitlaris Britton, Stern & Poggenb. USH 4456. Figure 42
Tri-zonocolporate. Ora lolongate 6.4 x 8.1 jjm. Exine 1.1 ]im thick.

Suboblate 23.8 x 29.5 [jm. Faintly reticulate.

Solatium -melongena L. SBGH 2/114. Figure 40
Tri-zonocolporate. Ora lolongate, forming a streak across the colpus

5.1 x 14.4 [j.m. Exine thickness 1.1 y.m. Prolate spheroidal 26.4 x 24.9
jim. Faintly foveolate.

Solanwm wrightii Nees. USH 4356. Figure 41
Tri-zonocolporate. Ora lalongate 6.0 x 4.9 um. Exine thickness 1.3

urn. Prolate spheroidal 21.4 x 19.8 um. Psilate.

Datura sziaveolens (Figure 38) is also studied (Erdtman 1952) but
based on observations it is found that the aperturate condition is colporate
in nature, but the species collected from Malaysia shows the brevicolpate

Figs. 37 - 40. 37, Datura metel. 38. Datura ettoveolens. 39. Capsionmannuu



REINWARDTIA

Figs. 41-49. 41. Solatium wrigktii. 42. Petunia, axittans. 43. Te
capensie. 44. Crossandra wndulaefolia. 45. Psexderanthemum retioulotum. 46. Berre
laevictadis, 47. Isotoma Umgiflora. 4B. Museaenda oblowga. 49. JiHsseito sarmevifo

condition. The ora present ia lalongate in the former species. Another
species found locally is Datura metel and this too shows a brevicolpate
condition. Apart from the two species studied locally, other species in
which their pollen grains are studied are Solanum alatwrn, SoUinwm
dulcamara, Solanum luteum, Solanum nigram (Erdtman 1952), Solatium
pseudocapsieum., and Solanum verbascifoKum (Nair 1965). In all cases,
there ia very little morphological variation and all of them show colporate
condition. The taxonomic significance of the pollen morphology in
Solanaceae is not so obvious because, sometimes genera belonging to the
same tribe or subtribe have similar pollen morphology, but in other cases,
it can be very different (Erdtman 1952).

The following table shows the species studied by Erdtman and Nair,
compared with the species presently studied.

Datura arborta (Chaubel
& Deodikar 1963)

RAO & LEONC: Tropioat pollen arai:

Aperturate conditions

Porate

Porate

Datura suaveoleua
(cited in Erdtman 1952)

Solatium peeudocapsicrtiTl
(Nair 1965)

Solanum vprbascifolitiin
(Nair 1B65)

Solanum ivrigkUi

Colporate

Brevicolpate

Colporate

B rev i col pate

Colpovate

Colporate

Colporate

Colporate

Colporate

Colporate

Colporatc

Colporate

Colporate

Colpomte

Colporate

35.8 X 35.9

37.!) x 39.3 j

37 X 40 j

40.4 X 33.1

41 X 45 jj.ni

23.6 x 21.4 jj.m

23.8 X 29.5 |im

30 x 26 :im

14.5 x 13 jtm

28.5 x 24.5 ^m

20 X 24 y,m

•26.4 X 24.9 y.m

17.5 X 14 .ij.m

SI x 18 :j.m

21.4 X 10.S (in

With reference to the table, the pollen grains of Datura stramonium
are smaller than the other three species. The species studied presently
(D. metel and D. sitaveolens) are brevicolpate, whilst other workers have
recorded colporate condition in certain species. Except for the size
Variations, the pollen of Solanum species generally show the colporate
condition.



REINWARDTIA [VOL. 1974] RAO LEONG: Tropical pollen gr 171

S C R O P H U L A K I A C E A E

Russelia sarmentosa Facq. USH 3293. Figure 49

Tri-zonocolporate. Ora lalongate 3.2 X 4.4 p.ni, Exine 0.9 y.m thick.
Oblate spheroidal 14.8 x 16.1 ij.m. Rugulate.

The pollen grains of members of this family have close affinitieH
with that of the genera in Solanaceae. But in this family, striate pattern
of exine, the n on-aperturate, or 4-aperturate conditions, and large size
of the grains are not so common as are encountered in members of
Solanaceae.

B I G N O N I A C E A E

Tecomatia, capeiisk Spach. USH 3322. Figure 43

Tri-zonocolpoidate. Exine thickness 1.6 ym. Oblate spheroidal
28.9 x 30.7 ;j.m. Retipilate.

ACANTHACLAE

Crossandm undulaefolia Salisb. USH 4360. Figure 44 Plate II, 6

Two colpate. Exine thickness 1.5 ;im. Exine thicker at equatorial
region, thinner at the poles. Perprolate 56.0 x 19.7 tim. Reticulate.

The grain is perprolate and 2-colpate. Normally, colpi are three
or more in number and perprolate condition is not frequently encountered
in pollen grains of other families. Pollen of this species studied earlier
also exhibit the perprolate condition, but larger in size (87 x 42 ;i.m)
(Erdtman 1952).

Pseuderwnthemum reticulatum Radlkf. USH 2402. Figure 45 Plate II, 7

Tri-zonocolporate. Ora lalongate, 5.2 x 3.8 ^m. Exine thickness
2.2 iim. Subprolate 39.8 x 32.5 y.m. Surface of the grain is scalloped and
this is best seen in the polar view. OL pattern punctate. The scalloped
shape of the pollen grain is quite unique, and characteristic.

The pollen of 3 other species described are P. em-datum, P. citatre-
casaii and P. malaecmise (Raj 1961). Essentially, they all show the
colporate condition.

R U B I A C E A E

Borreria laevicavMs Ridley. USH 5037. Figure 46

Zonoporate. Four to six pores are found in one pollen grain-
Pores slightly elongated. Pore diameter 1.8 jim. Pore margin 1.4 v-m-
Interpora! distance 10.6 ^m. Exine 1.4 ?.m thick. Prolate spheroidal
33.5 x 32.S ;j.m. Foveolate.

Mitssaenda oblonya King. USH 1790. Figure 48

4-Konoporate. Pore diameter 2.2 [im. Pore margin 0.8 im. Interporal
distance 7.6 7.111. Exine thickness 1 ;J.m. Oblate spheroidal 15.1 X 17 urn.
Foveolate.

Warseeiviezia eocciuea Klotzseh. USH 507. Figure 21
Tri-zonoporate. Pore diameter 2 {j.m. Pore margin 1.1 ;j.m. Exine

thickness 1.2 ;j.m. Prolate spheroidal 16.5 X 15.2 -jun. Excrescences
granulose. Retipilate.

Pollen morphological variations are seen with regard to shape, OL
pattern, aperturate condition and size.

CAMPANULACEAE

Isotonu), longiflora Presl. USH 3026. Figure 47 Plate 11, 2

Tri-zonocolpate. Exine thickness 2.2 y.m. Oblate spheroidal 35.3 x
37.7 j.m. Reticulate. Lumina in the polar region are very small, and as
they reach the equatorial region, they are very prominent.

Variations in pollen morphology are seen in this family. Three other
species studied are Isotoma (Lobelia) anceps var. minor: 29 x 19 ;im;
L.gloria-montis: 5 8 x 3 5 ;jm (Erdtman 1952) and L. nicotianaefolia:
2 6 x 2 3 vxa (Chaubal & Deodikar 1963). The aperturate condition in
the first two species listed is colporoidate. The latter has pollen grains
of the colpate condition. Size variations are indicated.

DISCUBSIOX

The pollen grains of plants studied either at the generic or species
level are given in the following table. For others the variations noticed
in the local plants are recorded and the data is compared with the earlier
accounts.

Studied at the genetic level

Imperata cylhidrica

Archoittophoenix alexandrae
twiwma picttttn
•aimlo. integrifolia.

Eichkornia crassipcs
Monochiria hastata

' The number of species studied pr

Studied at the
Aechmea Unctoria (2)"
Boerhaavia diffasa (14)
Portuluca grandiflora (1)
Braesica oleracea (2)
Moringa oleijera (1)
Weinmaimia blumei (1)
Ixonanlhes reticalata (2)
JabrQpha, podagriua (1)

level

sly is sho ithin the bracket.



172

Studied at the generic lev
Zephyrantkes alba
Geostackys rupestris
Andira surinamensis
Etaeedendron qaadrungulaUin
Gonocaryum Morale
Cerbera odollam
Roupellia grata
Tkevetia peruviana
Brunfelsia species (B. cafycin

I I A [VOL.

S t u d i e d a t t h e species level
Impatient griff ithii (S)
Wormia suffruticosa (1)
Rhododendron leucobotrys (4)
Styrax bemoin (2)
Raiiwolfia. vomitaria (1)
Ekretia microphylla (4)
Clerodendron specinsissimttnt (2)
Datura species (D. metel &

D. suaveolens) (2)
Solaiium species (S. melongena &

S. wrigktii) (6)
Fseuderantkemam reticulatum (3)
Isotoma longifloTa (3)

RAO & LEONG: TropieaJ jwMsn grains

Petunia axUlaris
Ruseelia sarmentosa
Tecotnariv* capeftsia
Borreria laevicaulie
Muaeaenda oblonga
Wareeewiaiia cocci-nea

Earlier, under the description of each species a brief comparison
is made about the pollen characters of the closely related genus or species.

In the palynological studies, the families studied may be broadly
divided into two groups, namely eurypalynous type, that is, their pollen
grains are characterised by a great array of pollen types and variations
in morphological characters like apertures, exine stratifications, OL
patterns, excrescences, etc.; the other group being the stenopalynous type
in which the pollen of different genera in a family show more or lesa
similar characters. The classification of the different families, presently
investigated, under the two groups is summarized below.

ies Stenopalynous families

Gramineac
Pontederiaceae
Zingiberaceae
Cruciferae

Euryp.

P a l
A r i
E n
Am

alynou.

raae
iccae
imeliac
arvllid

Mo ngaceae

Euphorbiacc

Apocynaceae
Verbenaceae

Acanthaceae

Celastrac
Balsainin:

Styracace

Scrovhulariaceae
Rubiaceae
Campanulaceae

Plate I. 1-i). 1. Impe
qiiitoewsia. A. Geostachys
1. Jatrophn porlagrica.

-ota eylindriea. 2. Pitca
rnpestris. B. Andira sili-i-
S. RhodtHlevdron U

i iittegrifolia. 3. Lejitochiton
ensis. 0. Ixonanthes reticulata.

•Iriie. !l. Nyetanth.es arbor-Math.

Among the species investigated the corporate (325!,), colpate (2i'/< j,
poi-ate (20^,), sulcate (12%) and non-aperturate (10%) grains are
found and the percentage of their occurrence is mentioned within the
brackets. Gramineae has pollen grains characterised by the presence of
a single pore. The non-aperturate condition is seen both in mono- and
dieots.

The thickest exine is seen in Boerliaavia diffusa (5.4 ;,m) and thinnest
in case of Geostachys i-iipestris (0.6 y.m). The smallest and largest grains



1 7 4 R E I N W A R D T I A [
V 0 L - f l

seen presently are those of Weinmannia bhimei (10.2 x 0.3 ;im) and
Geostachys rupestris (diameter 87.2 y.m) respectively.

Pollen characters are generally very specific and said to be genetically
controlled (Wodehouse 1935). The external characters of the pollen,
for example the diameter, size, number of pores or furrows sculpture of
exine are somewhat variable at least in some plants as a result of the

RAO & LEOKO: Trapical polkn gr 176

Plat* II 1-S 1 Rauwolfia vomitoria. 2. leotamu Kingiflora. 3. Ckrodendi
speciosis'sinmm 4 Brunfelsia calycina. 5. Datura *uavcolew>. S. Crosiandra tindul

folia 7 Pseuderanthemum retittulatum. S. Ehretia microphyUa.

climatic effects on the pollen forming plants or directly upon the process
of pollen formation (Kurtz & Liverman 1958). High temperature,
repeated cooling of flower buds, variation in the moisture contents of
the soil or arid condition may effect the size of the grains, or number
uf germ pores on them (Kurtz & Liverman 1%8). Polyploid condition
is another factor that causes variations in pollen size and viability within
a given species. Among the cultivated plants such variations are common.
Some workers claim that the position of the flower on the plant, the
size of the anther, and the time of flowering also influence the pollen size.
Among1 the plants presently studied such variations are noticed in the
following genera: Portulaca, Ixonanthes, Nyctanthes, Datura and
Cros&andra. The exine pattern of the pollen of Nyctanthes arbor-trigtis
found locally is retipilate and do not possess any excrescence, in contrast
to that found in Thailand which have a reticulate pattern with more or
less piloid excrescences (Erdtman 1952).

HEFERENOES

CHAUHAI, P. D. & FIEODIKAB, G. B. (Iflfi3>. Pollen grains of poisonous plants.
I. Poisonous pollen in honey samples from Western Ghats (India). hi Grana
PalynoL i: 393-397.

CUANWRLL, L. M. (1953). New Zealand poliea stttdies. The tnonneotytedons. Harvard
University Press, Cambridge, Mass.

EEDTMAU, G. (1943). An introduction !u pollen anali/sh. Chronic Botanies Company,
Walttam, Mass.

ERDTMAN, G. (1952). Pollen morphology and plant taxonomy. Almqvist & Wikscll,
Stockholm.

EKDTMAN, Ci. (1957J. Pollen and spore nunjiholoBy/plant taxotiamy. Almiivist &
Wiksell, Stockholm.

KHDTMAM, G. (I960). The aeetolysis method. A revised description. In Svensk.
Bot. Tidskr. 54: E61-564.

EHDTMAN, G., BEHGLUNEL, B. & PRAGLOWSHI, J . (1961). An introduction to n

Scandinavian pollen flora. Alraqvist & Wiksell, Stockholm.
PAF.OHI, K. (19B6). Recent trends in palynology. In Bot. Rev. 22: 639-684.
FARCHT, K. & IVERBEN, J. (1950). Textbook of mode™ pollen analysis. Munksgaard,

Copenhagen.
PAECRI, K. & IVERSsr.-, J. (1964). Textbook of pollen analysis. Munksgaard, Copen-

hagen.
HYDE, H. A. & ADAMS, A. F. (1958). An atlas of airborne pollen grant*. Macmillan

& Co., London.
KEN'C. H. (1S70). Size and affinities of the ftora of the Malay Peninsula. la Jour.

Trap. Geog. 3 1 : 43 - 66.
KURTZ, B. B. & LTVEBMAN, J. L. |19BS). Some effects of temperature on pollen

characters, hi Bull. Torrey Bot. CI. 85: 136-138.
NAIR, P.K.K. (1965). Pollen grains of Western Himalayan plants. Asia, Bombay.



REINWARDTIA

Pollen morphology in the Nytaginaceae . In Grana Palynol

Morphological studies in the Acanthaceae. In Grana Palynol

(1970).Studies on singapura pollen. in pacific Sci

(1W0).

HEMNWABDTIA
Published by Herbarium Bogoriense — LBN, Bogor

Vol. %, Port 1, pp. 177 — 182 (1974)

ON SOME COLOURLESS FLAGELLATES
FROM JAVA AND BRASIL

B. V. SKVORTZOV
Instituto de Botanica, Sao Panic, Brazil

ABSTRACT

Two new monotypie genera (Kaakimonas bogoTiensis and Hoehne-
mastix saepatilensiB) and aiK other new species (four in Tetramitus and
two in Balliamonas) of colourless flagellates are described based on
samples collected in Bogor and Sao Paulo.

ABSTRAK

15ua ntnYgn inonotipe {KtzaKi'itioiiaB hoffOTicnsts dan nochyiswtastix
xaopatilpnsis) dan enam jenis (empat dalam Tetramitus — antara lain
Tetrumiius indowe&iae dan dua dalam Baltiatnwiasi flaeellata tak

contoh yang dikumpulkan di Bogor (Jawa) dan Sao Paulo (Brasilia).

The colourless flagellates described in the present note have been
cultivated and studied in the Instituto de Botanica, Sao Paulo. They
were isolated from samples collected by Dr. M. Kizaki in Bogor (Java)
in 1971 and by the present author in Sao Paulo, Brasil. All illustrations
presented were made from living specimens and all type specimens studied
are preserved in the Cryptogamic Herbarium of the IiiKtituto de Botanica,
Sao Paulo, Brasil.

The eight species described are distributed among four genera which
can be distinguished as follows.

1. Cells 3 times broader than its length, with 2

2. Cells fusiform, with 2 anterior and one poster]

3. Cells elongate to fusiform, with 3 s

4. Colls elongate ovate to fusiform, wit!

•imming flagelks.

terior flageiles.
BaUiamantui Sltv

flagelles of different length.
Haohnen

• flagelles of diffe
Perty

Kizakimonas Skvortzov, gen. nov.

Fam. Ampbimonadaceae, Ord. Protomastiginae.
Monada solitaria, libere jiatantes, compresso-lanceolata, cum antico

et postico oppositis convexie, 3 plo latius quam longius; membrana



i C O N T E N T S

P a g e

H A T T I N K , T. A. A revision of Malesian Caesalpinia, i n c l u d i n g
, Mezoneuroji ( L e g u m m o s a e - C a e s a l p i n i a c e a e ) 1

• J O N E S , H . G < Orchidaceae n a v a e vel m i n u s cogtlitae . . . . . . . 7 1

K E N G , H. Rediscovery of Cheilotheca malayana a n d t h e i d e n t i t y of .
Cheilotheca, Audresia and Mo.notropastmm (Ericaceae- .
M o n o t r o p o i d e a e ) 7 7

K O S T E R M A N S , A . J . G . H . A m o n o g r a p h o f t h e genus- N.eoanna-

•momum L i o u H o 8 5

M a t e r i a l s f o r a r e v i s i o n o f L a u r a c e a e I V . . . . . 9 7

r? A new Bornean species .of Mammea , . 117

Triadodapkne, a. new Jauraceous genua from Borneo . 119

— A monograph of Caryodaphnopsis A. Shaw . ., . . . 123

LARSEN, K. & LAKSEN, S. K. A new Amorphophallus from Thailand ' 139

NAYAK, M. P. A revision of Phtkiandra (Melastomataceae) . . 143

RAO, A. N. £ LEONG, F. L. Pollen morphology of certain tropical ,
• plants . . . . . . . - . • 153

SKVORTZOV, B. V. On some colourless flagellates from Java and Brasil 177

Distributor
BIBLIOTHECA BOGORIENSIS,

JALAN RAYA JUANDA 20,
EOGOK, INDONESIA

by ARCHIPEL


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