Gillespie1994 - Polen - Compressed

Pollen Morphology and Phylogeny of the Tribe Plukenetieae (Euphorbiaceae)
Author(s): Lynn J. Gillespie

Source: Annals of the Missouri Botanical Garden, Vol. 81, No. 2 (1994), pp. 317-348
Published by: Missouri Botanical Garden Press
Stable URL: http://www.jstor.org/stable/2992101 .
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POLLEN MORPHOLOGY AND Lynn J. Gillespie2
PHYLOGENYOF THE TRIBE
PLUKENETIEAE
(EUPHORBIACEAE)1
ABSTRACT
A scanning electron microscopy and light microscopy survey of pollen morphology in the Plukenetieae (Euphor-
biaceae) was undertaken to help elucidate phylogenetic relationships within the tribe. Pollen is medium to large,
spheroidal to suboblate, and tricolpate, inaperturate, or with poorly defined apertures. Subtribe Plukenetiinae is
characterized by tricolpate pollen with uneven-margined colpi and a perforate to reticulate tectum. Pollen evidence
supports a division between genera having an aborescent habit (Angostyles, Astrococcus, and Haematostemon) and
those with a scandent habit (Plukenetia and Romanoa). The synonymy of the monotypic genera Vigia and Eleuthero-
stigma with Plukenetia is also supported. Subtribe Tragiinae is exceptionally diverse in pollen morphology. Aperture
condition ranges from tricolpate, the plesiomorphic and most common state, to weakly aperturate and inaperturate;
islands, fragments, or strands of sexine are usually present on the apertural membrane, and aperture margins are
uneven and often indistinct. Exine sculpture is punctate, foveolate, reticulate, rugulate, or baculate. The large genus
Tragia includes seven distinct pollen types, with most sections (e.g., Bia, Ctenomeria, Leptobotrys, Tragia, and
Zuckertia, and also subgenus Mauroya) characterized by a uniform and unique pollen morphology, supporting the
sectional classification of Tragia. The other Tragiinae genera have pollen distinct from Tragia, with the exception
of Tragiella, which closely resembles sections Tagira and Lassia. Pollen evidence supports Cnesmone and Megis-
tostigma as sister taxa, and suggests a close relationshipwith Pachystylidium. Acidoton includes two different pollen
types; the inaperturatetype closely resembles pollen of Platygyna, suggesting that Acidoton may not be monophyletic
and the tricolpate species perhaps represents a distinct genus. Pollen, together with floral morphological evidence,
supports the hypothesis of section Zuckertia as a plesiomorphic member of Tragia, and suggests that Tragia is
paraphyletic and that the smaller Tragiinae genera are derived from Tragia.
The Plukenetieae belong to the Acalyphoideae, scription of genera and infratribal phylogenetic re-
the largest and least understood of the five eu- lationships are the principal systematic problems
phorbiaceous subfamilies. The tribe includes 13 in the Plukenetieae.
genera distributed worldwide in tropical and warm Pollen morphology has been invaluable in the
temperate regions. Many species are twining vines systematics of the Euphorbiaceae (Punt, 1962;
or lianas, both unusual habits in the family; other K6hler, 1965). Light microscopic (LM) observa-
species are erect herbs, shrubs, or rarely small tions by Punt (1962), in his pollen survey of the
trees. Although flowers are small and apetalous, Euphorbiaceae, revealed a diversity of pollen types
floral morphology is diverse, particularly the style among species belonging to the Plukenetieae. The
and androecium. Another uncommon feature is the present study of pollen morphology of the Pluke-
presence of stinging hairs in many species. Tragia netieae based on scanning electron microscopy
L. is the largest genus, with more than 125 species, (SEM) and LM was initiated to help resolve prob-
some of which are commonly known as nose burns. lems of generic circumscription and elucidate phy-
All other genera have fewer than 17 species each; logenetic relationships. The study is the first part
many are monotypic or with few species. Circum- of a larger project concerning the evolutionary
' I offer my warmest thanks to G. Webster for support and encouragement. Much of the work was undertaken
while I was working as a graduate student in his laboratory at the University of California, Davis. The Facility for
Advanced Instrumentation, University of California, Davis, was extremely helpful in SEM instruction and in solving
problems encountered in the SEM work. I thank J. Nowicke for offering lab facilities and photographic supplies, and
C. Hotton for helpful comments on the manuscript. The curators at A, DAV, MO, NY, P, UC, and US are gratefully
acknowledged for the loan of specimens and for removing or allowingpollen to be removed. I thank several individuals
for their SEM photographs, which form part of G. Webster's Systematics Laboratory pollen research collection at
the Botany Dept., UC Davis. S. Lynch was responsible for Figures 23, 24, 29, 39, and 42; J. Ward was responsible
for Figures 19 and 54. All remaining SEM work and photography were performed by the author.
2
Botany Department, Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ANN. MISSOURIBOT. GARD. 81: 317-348. 1994.

318 Annals of the
Missouri Botanical Garden
history of the Plukenetieae, which will include cla- Croizat (1941) and Airy Shaw (1969); of the two
distic analyses based on floral, vegetative, and other species treated by Pax and Hoffmann, M. malac-
characters, studies of trends in character evolution, cense Hook. f. was considered a species of
and an attempt at correlating the unusual and often Sphaerostylis Baill. and M. peltatum (J. J. Sm.)
bizarre diversity in floral morphology with polli- Croizatas the monotypic Clavistylus J. J. Sm. The
nation biology. Determining plesiomorphic char- genus Ramelia Baill. has since been reduced to
acter states and the direction of evolutionary trends synonymy under Bocquillonia Baill. (Acalyphoi-
will also be of importance in phylogenetic studies deae, tribe Alchornieae) and thus excluded from
of other genera for which the Plukenetieae or its the Plukenetieae (Airy Shaw, 1968, 1974), while
members have been considered as outgroup(s), such Megalostylis is now considered to be a synonym
as Dalechampia L. (Armbruster, 1994) and Om- of Dalechampia (Webster & Armbruster, 1991).
phalea L. (Gillespie, 1988). In his most recent classification, Webster (1994)
There have been many additions and taxonomic treats Dalechampia as a subtribe, the Dalecham-
changes in the Plukenetieae since it was first rec- piinae, of the Plukenetieae, rather than as a distinct
ognized as a distinct group by Bentham (1880, as but related tribe (as in Webster, 1975). Although
subtribe "Plukenetieae"). The circumscription of the genus will not be extensively treated in the
the tribe and its genera recognized closely follows present paper, its pollen morphology will be dis-
that of Webster (1975, 1994) in his classification cussed in relation to the Plukenetieae.
of the Euphorbiaceae. Two exceptions are recog- In the first attempt at an infratribalclassification
nition of the genera Tragiella Pax & K. Hoffm. and consideration of relationships, Pax & Hoff-
(following Radcliffe-Smith, 1982, 1987) and mann (1919) divided the 19 genera of their "Plu-
Pachystylidium Pax & K. Hoffm. (following Airy kenetiinae" into four informal groups, Plukenetii-
Shaw, 1969, 1975), both originally described as formes, Astrococciformes, Tragiiformes, and
species of Tragia. In addition, several taxonomic Sphaerostyliformes, based primarily on stamen
and nomenclatural changes have taken place since number and style shape. Webster (1975) created
Webster's 1975 synopsis. The monotypic genera the first formal subtribal classification upon de-
Vigia Vellozo (which has priority over the more scribing the Tragiinae. The Tragiinae are char-
commonly used generic name Fragariopsis A. St. acterized by presence of stinging hairs, a trilocular
Hil.) and Eleutherostigma Pax & K. Hoffm. have ovary, and absence of foliar glands; species are
been synonymized under Plukenetia (Gillespie, found in all habitats but are particularly diverse in
1993). Adrien Jussieu's genus Anabaena has been dry areas. In contrast, the Plukenetiinae are char-
twice renamed, as Romanoa by Trevisan (1848) acterized by absence of stinging hairs, foliar glands
and as Anabaenella by Pax & Hoffmann (1919), typically present and an ovary that is usually
due to similarity with its namesake cyanobacteria. 4-locular or less often trilocular, and are usually
Although the cyanobacteria were originally de- found in wet habitats. Style morphology has been
scribed as Anabaina Bory, the more commonly used extensively in infratribalclassification and ge-
used name of Anabaena Bory was recently con- neric delimitation. With the exception of the genus
served against Anabaena Adr. Juss. (ICBN, Greu- Tragia, there has been an unusual radiation in
ter, 1988: 112). Romanoa, the name rediscovered style morphology in the Plukenetieae. Within sub-
by Punt (1962) and Radcliffe-Smith (1980), thus tribe Tragiinae, many of the genera are charac-
becomes the valid name for the genus. terized and differentiated from Tragia on the basis
The most recent monograph of the Plukenetieae of unusually shaped massive styles (e.g., Sphaero-
(as subtribe Plukenetiinae of tribe Acalypheae) was stylis, Megistostigma, Cnesmone Blume, and Tra-
by Pax & Hoffmann (1919; with additions and giella). Pax & Hoffmann (1919, 1931) based their
changes, 1924, 1931). The authors treated a num- sectional classification of Plukenetia on style mor-
ber of segregate genera in addition to Tragiella phology.
and Pachystylidium that were not recognized by Since Tragia is a large and diverse genus it is
Webster (1975); these are Gitara Pax & K. Hoffm., necessary to consider an infrageneric classification.
considered synonymous with Acidoton Sw. (Web- Eight sections are recognized in the present study.
ster, 1967), and Tetracarpidium Pax (also known The sectional classification of Pax & Hoffmann
as Angostylidium (Muell. Arg.) Pax & K. Hoffm.), (1919, 1931) is followed with the following excep-
Apodandra Pax & K. Hoffm. and Pterococcus tions. Section Leptobotrys (Baill.) Muell. Arg. is
Hassk., all presently treated under Plukenetia L. considered distinct from section Tragia, whereas
(Gillespie, 1993). A second difference is Webster's sections Leucandra (Klotzsch) Muell. Arg. and Ra-
recognition of Megistostigma Hook. f. following tiga Muell. Arg. are treated as part of section

Volume 81, Number 2 Gillespie 319
1994 TribePlukenetieae
Tragia (following Miller & Webster, 1967, and considered part of section Tragia by Pax & Hoff-
Mligura de Romero & Gutierrez de Sanguinetti, mann (1919), and to doubt the taxonomic validity
1989). Also considered is L6andri's subgenus Mau- of section Leucandra (Klotzsch) Muell. Arg.
roya, which includes a single Madagascan species.
Many of the sections have in the past been rec-
METHODS
ognized as distinct genera (e.g., Bia Klotzsch,
Ctenomeria Harv., Lassia Baill., Leptobotrys Baill., Flowers were removed from herbarium speci-
Leucandra Klotzsch, and Zuckertia Baill.). mens, of which determinations were verified, and
Previous pollen morphological studies of the Plu- rehydrated in 3% Aerosol-OT for 3-4 days. Pollen
kenetieae have been based only on LM (Erdtman, grains were isolated and mounted in Hoyer's me-
1952; Punt, 1962; Miller & Webster, 1967), with dium (Radford et al., 1974) for examination under
the exception of studies dealing only briefly with LM. The method of Lynch & Webster (1975) was
pollen morphology (e.g., Gillespie, 1988; M6lgura followed for SEM studies. Pollen grains were de-
de Romero & Gutierrez de Sanguinetti, 1989). The hydrated to 100% ethanol, to 100% amyl acetate,
most extensive survey was that of Punt (1962), and then critical-point dried. Following sputter
who examined 17 genera (equivalent to 12 in the coating with gold or a gold/palladium mixture,
classification followed here) in his pollen survey of grains were examined and photographed in an ISI
the Euphorbiaceae. Pollen types within the tribe DS130 (equipped with a LaB6 filament), Hitachi
fell into two of Punt's 17 main groups, the Plu- S800, or Cambridge 250 scanning electron mi-
kenetia configuration and the Cnesmosa (= Cnes- croscope. All pollen was prepared in the above
mone) configuration. The Plukenetia configuration manner unless specified otherwise. In several cases
is characterized by oblate-spheroidal to oblate, tri- pollen was acetolyzed (following the method of
colpate or triporate grains with broad apertures Erdtman, 1952) then mounted directly on a stub
having a "ruptured membrane." Of the two types and sputter coated for examination under SEM.
within this main group, the Plukenetia type is Voucher microscope slides and electron micro-
tricolpate and includes all examined species of the graphs are deposited at the Systematics Labora-
Plukenetiinae, African Tragiinae, and most New tory, Botany Department, University of California,
World Tragia spp. The Pachystylidium type is Davis. Voucher LM slides of acetolyzed pollen are
triporate and includes only that genus. The Cnes- deposited at the Palynology Laboratory, Botany
mosa configuration is characterized as inaperturate Department, Smithsonian Institution, Washington,
and lacking a crotonoid exine; three of its five types D.C.
are composed of species of the Plukenetieae. The Descriptions of pollen grains are based on ob-
Cnesmosa type of pollen with a psilate exine in- servations under LM and SEM. Measurements were
cludes Acidoton, Megistostigma, and Cnesmone. made under LM on 15 grains mounted in Hoyer's
The Tragia fallax type of pollen is described as medium. The polar axis (P), equatorial axis (E),
having an "intectate pilate exine" and consists of and the polar to equatorial axis ratio (P/E) are
Tragia sect. Bia (Klotzsch) Muell. Arg. The Platy- given in the generic descriptions as the range of
gyne (= Platygyna) type of pollen with an exine mean values of collections examined (refer to Table
that is tectate and "intra-reticulate" (defined by 1 for measurements of individual collections). In
Punt as "columellae inside the tectum form[ing] the case of inaperturate grains that are ellipsoidal
a network") consists only of species of Platygyna in shape, the shortest axis (S) and longest axis (L)
Mercier Punt concluded that pollen morphology are given. Exine thickness of aperturate grains was
supported the distinctness of Tragia sect. Bia, the measured at mid mesocolpium in polar view. Ter-
close relationship of Haematostemon (Muell. Arg.) mmology used follows that of Erdtman (1 952, 1966)
Pax & K. Hoffm., Angostyles Benth., and Astro- and Walker & Doyle (1975). In the present paper
coccus Benth., and the observation that Apodan- the term scabrate is restricted to the description
dra, Romanoa, Fragariopsis (= Vigia), Pterococ- of a surface having a covering of microprojections
cus, and Angostylidium cannot be easily that are irregular in size, shape, and distribution
distinguished from Plukenetia. Pollen evidence also (e.g., Figs. 27, 48, 69). The term microverrucae
supported Croizat's (1941) circumscription of (e.g., Figs. 9, 38, 41, 43) is introduced to refer
Sphaerostylis and Megistostigma. Miller & Web- to microprojections that are more regular in size,
ster (1967) examined pollen of Tragia species from wider than high, and rounded in shape but not
the United States; evidence from their study of constricted at the base (as distinguished from con-
pollen and floral morphology led them to revive ical spinules and constricted-based microgemmae).
section Leptobotrys (Baill.) Muell. Arg., which was Twelve of the 13 genera in the Plukenetieae

TABLE 1. Species of Plukenetieae examined with voucher information, geographical location, pollen size dimensions (min
equatorial axis (E Axis), all in ,.m, and pollen shape given as ratio of polar to equatorial axes (P/E). Inaperturate pollen is i
and longest axis (under E Axis) are given if pollen shape is ellipsoid, otherwise a single diameter is given if spheroidal. Pollen w
by a "c"; shortest and longest axes are given. Measurements of acetolyzed pollen are preceded by an "a."
Species Collection Location

Plukenetiinae
Angostyles longifolia Benth. Spruce 2282, NY Brazil
Astrococcus cornutus Benth. Liesner 8693, NY Venezuela
A. cornutus Benth. Liesner 8693, NY Venezuela
Eleutherostigma lehmannianum Pax & K. Hoffm. Cazalet & Pennington 5089, UC Ecuador
Haematostemon coriaceus (Baill.) Pax & K. Hoffm. Wurdack & Adderley 43210, NY Venezuela
H. guianensis Sandw. Fanshawe 2869, US Guyana
Plukenetia africana Sond. Pope et al. 834, MO Botswana
P. africana Sond. Wild 5062, MO Botswana
P. brachybotrya Muell. Arg. Vargas 18799, US Peru
P. conophora Muell. Arg. Zenker 3394, US Cameroon
P. conophora Muell. Arg. Zenker 3394, US Cameroon
P. corniculata Smith Koorders 41720, UC Indonesia
P. loretensis Ule Fosberg 29094, MO Peru
P. loretensis Ule Maguire & Politi 27371, US Venezuela
P. penninervia Muell. Arg. Werff & Wingfield 3173, DAV Venezuela
P. penninervia Muell. Arg. Standley 56708, A Honduras
P. polyadenia Muell. Arg. Lindeman 6134, DAV Surinam
P. madagascariensis L6andri Morat 4893, P Madagascar
P. multiglandulosa Jabl. Cowan & Wurdack 31400, US Venezuela
P. stipellata L. J. Gillespie Gillespie 418, DAV Costa Rica
P. supraglandulosa L. J. Gillespie Cowan 38204, US French Guiana
P. verrucosa Smith Prance et al. 11255, DAV Brazil
P. volubilis L. Asplund 14129, US Peru

TABLE 1. Continuled.

Romanoa tamnoides (Adr. Juss.) A. Radcliffe-Smith Webster et al. 25436, DAV Brazil
Vigia serrata Vell. Bradem et al. 8376, DAV Brazil
V serrata Vell. Hoehne 29250, A Brazil
Tragiinae
Acidoton nicaraguensis (Hemsley) Webster Ortiz 1104, DAV Nicaragua

A. nicaraguensis (Hemsley) Webster Steyermark & Davidse 116239, DAV Venezuela
A. urens Swartz Proctor 36826, MO Jamaica
A. microphyllus Urb. Leonard 5248, US Haiti
Cnesmone anisosepala (Merr. & Chun) Croiz. Lau 141, UC China
C. javanica Blume Morse 567, NY China
C. philippinensis (Merr.) Airy Shaw Ramos & Edano 47087, UC Philippines
C. tonkinensis (Gagnep.) Croiz. Petelot 6521, A Vietnam
Megistostigma cordata Merr. Ramos 17591, US Philippines
M. malaccense Hook. f. Rahmat si Toroes 1389, A Indonesia
M. malaccense Hook. f. Burkhill & Haniff 15589, MO Malaysia
Pachystylidium hirsutum (Blume) Pax & K. Hoffm. Clemens 1748, UC Philippines
P. hirsutum (Blume) Pax & K. Hoffm. Ramos & Edano 49201, UC Philippines
Platygyna hexandra (Jacq.) Muell. Arg. Howard et al. 81, A Cuba
P. hexandra (Jacq.) Muell. Arg. Jack 7146, US Cuba
P. leonis Alain Leon 12176, DAV Cuba
P. parvifolia Alain Schafer 1427, NY Cuba
Tragia adenanthera Baill. Tanner 672, UC Tanzania
T bailloniana Muell. Arg. Cowan 2692, DAV Mexico

TABLE I Continued.
T capensis Thunb. Ecklon & Zeyher s.n., US 1170932 South Africa

T capensis Thunb. Kuntze s-n., NY South Africa
T chlorocaulon Baill. Eiten & Eiten 4285, US Brazil
T. cordifolia Vahl Webster s.n., DAV Kenya
T hispida Willd. Nicolson 2991, US India
T involucrata L. Nicolson et al. HFP173, US India
T. ivohibeensis Leandri Humbert 3387, DAV Madagascar
T lessertiana (Baill.) Muell. Arg. Webster 24119, DAV Surinam
T mexicana Muell. Arg. Tuerckheim 7664, US Guatemala
T novae-hollandiae Muell. Arg. Dovey-B56, UC Australia
T pacifica McVaugh McVaugh 21006, DAV Mexico
T peltata Klotzsch dos Santos 1551, DAV Brazil
T polyandra Vell. Bresolin 629, US Brazil
T ramosa Torr. Ferris & Bacigalupui 8136, DAV U.S.A.
T scandens (Baill.) Muell. Arg. Humbert 13741, P Madagascar
T sellowiana (Klotzsch) Muell. Arg. Webster 25463, DAV Brazil
T smaliji Shinners Curtiss s.n., US U.S.A.
T tristis Muell. Arg. Anderson 9117, DAV Brazil
T urens Small Norris 759, DAV U.S.A.
T volubilis L. Webster & Proctor 5325, DAV Jamaica
Tragiella natalensis (Sond.) Pax & K. Hoffm. Mearns 295, NY Kenya
T natalensis (Sond.) Pax & K. Hoffm. Mearns 295, NY Kenya

1994 Tribe Plukenetieae
2Eve
~~~ ~~~ 4~~
ORA ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~-' ,.
Y, Jr~~~~~~~~~~~V'
~~~~~~~~~~~ 4'~~~~~~~~~~~~~~~~~~~~~~~~~~T
/ Lw~~~~~~~~~~~~~~~A
~~~.~ P
4.~~~~~~~~~p
FIGURES1-6. Scanning electron micrographs of pollen of Angostyles and Astrococcus (subtribe Plukenetiinae).
1-3. Angostyles longifolia.-1. Polar view.-2. Close-up of colpus.-3. Exine sculpture. 4-6. Astrococcus cor-
nutus.-4. Close-up of colpus.-5. Polar view.-6. Equatorial view. Scale bar: = 10 ;1m in Figs. 1, 5, 6; = 5 gm
in Fig. 2; = 2 Am in Figs. 3, 4.
3 Voucher informationis in Table 1 unless given in the caption.

324 Annals of the
Iq
'W
hi 11i. ;~/ q~~4
I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i it -f
FIGURES 7-13.3 Scanning electron micrographs of pollen of Eleutherostigma and Haematostemon (subtribe
Plukenetiinae). 7-8. Eleutherostigma lehmannianum.-7. Exine structure of mesocolpium of fragmented grain.-
8. Oblique view. 9-13. Haematostemon coriaceus.-9. Close-up of colpus in equatorial view lacking a continuous

and a total of 55 species were examined (Table 1). angulaperturate; colpus broad with ends often in-
Pollen of Sphaerostylis, a small genus of vines distinct, margins uneven and jagged; exine tectate-
endemic to Madagascar, and one section of Tragia perforate, 2-2.5 Am thick, somewhat thinner at
endemic to Madagascar, Agirta Baill., was not colpus margin; tectum foveolate, surface smooth.
available.
Haematostemon (Figs. 7-13). Two species
RESULTS SUBTRIBE PLUKENETIINAE of shrubs or small trees in Guyana and Amazonas,
Venezuela, both examined.
Angostyles (Figs. 1-3). A monotypic genus
Pollen suboblate to oblate-spheroidal (P/E =
of small trees known only from the Rio Negro
0.86-0.89), 35-44 Am P x 40.5-49.5 Aim E,
region of Amazonian Brazil.
tricolpate; amb subcircular; colpus narrow, some-
Pollen suboblate (P/E = 0.87), 54 Am P x 62
times covered with an unbroken sexinous mem-
Atm E, tricolpate; amb subcircular to obscurely
brane, margins uneven; exine tectate-perforate,
obtuse-triangular; colpus narrow and short, mar-
1.5-2 Aim thick, becoming thicker, 3-3.5 Aim, at
gins uneven; exine tectate-perforate, 1-1.5 gm
colpus margin, with upper and lower exine layers
thick, uniformly thickened; tectum very finely fo-
separating forming an elongate chamber in the
veolate-rugulate, fragmented and irregularly finely
vicinity of the colpus; tectum very finely foveolate-
gemmate at colpus margin, rugae with evenly spaced
rugulate, becoming fragmented and irregularly
microverrucae, intervening perforations narrow,
gemmate at colpus margin, rugae with (1-)2(-3)
sinuous to sometimes small and circular.
rows of evenly spaced microverrucae, intervening
Astrococcus (Figs. 4-6). A monotypic genus perforations narrow, sinuous to small and circular;
of shrubs or small trees in the upper Rio Negro apertural sexine, when present, densely and irreg-
region of Venezuela and Brazil. ularly scabrate.
Pollen suboblate to oblate-spheroidal (P/E - Grains within a single SEM preparation have
0.88), 48 gm P X 54.5 ,um E, tricolpate; amb apertures that appear to have a sexine that is
subcircular to obscurely obtuse-triangular; colpus continuous (Figs. 9, 10) or very fragmented (or
very narrow and short, sometimes covered with an sometimes absent) (Figs. 7, 8, 11, 12) over the
unbroken sexinous membrane, margins uneven; ex- apertures. This difference may perhaps be due to
ine tectate-perforate, 1.5-2 ,m thick, becoming degree of rehydration or tolerance to treatment
thicker, ca. 2.5-3 ,im, and distinctly raised at with acetone or sonication, resulting in some grains
colpus margin, with upper and lower exine layers having a ruptured or degraded apertural sexine.
separating forming an elongate chamber in the
Plukenetia (Figs. 14-24). A genus of 16
vicinity of the colpus; tectum very finely foveolate-
species of twining vines and lianas distributed pan-
rugulate, fragmented and irregularly, finely gem-
tropically with one species in Asia, four in Africa
mate at colpus margin, rugae with 2(-3) rows of
and Madagascar, and 11 in the Neotropics (note
evenly spaced microverrucae, intervening perfo-
that the two species previously treated as species
rations very narrow, sinuous to small and circular
of Eleutherostigma and Vigia are included in the
particularly near the colpus.
species count but are described separately). The
A continuous unbroken sexine may sometimes
13 species examined may be divided into two pollen
be present over the apertures as in pollen of Hae-
types based on tectum morphology.
matostemon (Figs. 1 1, 12; refer to description and
Type 1 (Figs. 14-19). Pollen suboblate to ob-
discussion under Haematostemon).
-late-spheroidal (P/E = 0.80-0.89), 35-56 ,um P
Eleutherostigma (= Plukenetia) (Figs. 7, x 42.5-64.5 ,um E, tricolpate; amb obtuse-tri-
8). A monotypic genus of lianas of premontane angular to subcircular, angulaperturate; colpus
forest in Colombia and Ecuador, its single species broad with margins very uneven and jagged; exine
now treated as a species of Plukenetia, P. leh- tectate-perforate, 1.5-4 jim thick; tectum foveo-
manniana (Pax & K. Hoffm.) Huft & L. J. Gil- late with foveolae sometimes becoming smaller at
lespie. aperture margin, surface smooth or scabrate (e.g.,
Pollen oblate-spheroidal (P/E = 0.89), 50.5 ,m P. stipellata).
P x 56.5 ,m E, tricolpate; amb obtuse-triangular, Of the seven species included in Type 1, the
apertural sexine. -10. Close-up of colpus of grain in Fig. 13; note lack of an apertural sexine. -11. Equatorial
view.- 12. Close-up of colpus of grain in Fig. 11; note presence of sexine covering aperture.- 13. Polar view of
grain lacking a distinct apertural sexine. Scale bar: = 10 Atmin Figs. 8, 11, 13; = 2 Atmin Figs. 7, 9, 10, 12.

326 Annalsof the
MissouriBotanicalGarden
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41P ~ ~ ~ ~ ~ "
s~~~ *v;. A: .1
-~~~~~~~~~~:1
FIGURES14- 19.3 Scanning electron micrographsof pollen of Plukenetia (pollen Type 1). 14-16. P. stipellata. -
14. Polar view.- 15. Equatorial view. -16. Exine sculpture.-17. Polar view of P. polyadenia. -18. Polar view
of P. africana (Wild 5062 MO).- 19. Equatorial view of acetolyzed grain of P. conophora. Scale bar: = 10 tim in
Figs. 14, 15, 17-19; = 2 Am in Fig. 16.

Volume81, Number2 Gillespie 327
4%~~~~~~~~~~~~~4
- .~ ~ ~ ~~~~~~~~~~~~~~~~~~A
- -~ ~ ~ ~ ~ ~ ~~~~~~~W
211
FIGURES20-24.3 Scanning electron micrographs of pollen of Plukenetia(pollen Type 2). 20-22. P. loretensis
(Maguire & Politi 27371 US). -20. Polar view.-2 1. Exine sculpture.-22. Equatorialview. 23-24. P. penninervia
(Standley 56708 A).-23. Oblique view.-24. Close-up of colpus and exine sculpture. Scale bar: = 10 Jim in Figs.
20, 22, 23; = 5 /Amin Figs. 21, 24.
New World species, Plukenetia polyadenia (Fig. Type 2 (Figs. 20-24). Pollen suboblate (P/E
17), P. stipellata (Figs. 14-16), and P. volubilis = 0.78-0.85), 32-55 Itm P x 41-66.5 /Am E,
have large pollen grains (57-69 ,um E) with an tricolpate; amb obtuse-triangular, angulaperturate;
amb usually obtuse-triangular and an exine 2-4 colpus broad with margins very uneven and jagged;
,.rm thick that becomes gradually thinner at the exine semitectate-reticulate, 2.5-4.5 Im thick; muri
colpus margin. The Old World species, P. africana usually crenate (i.e., tranversely ridged), becoming
(Fig. 18), P. conophora (Fig. 19), P. corniculata, fragmented and sometimes finely geminate at col-
and P. madagascariensis have medium-sized pol- pus margin, lumina often smaller near colpus mar-
len grains (41-50 ,um E) with a subcircular amb gin.
and an exine 1.5-2.5 Ium that is mostly uniformly Of the six species included in Type 2, Plukenetia
thick or sometimes thickened at the colpus margin. brachybotrya and P. loretensis (Figs. 20-22) have

328 Annalsof the
-N AA-f-*-
I 'S ~~~~~~~~~0
4 .
1 I -, -R.
FIGUREs 25-29.3 Scanning electron micrographs of pollen of Romanoa and Vigia (subtribe Plukenetiinae). 25-
27. Romanoa tamnoides. -25. Polar view.-26. Close-up of colpus of grain less expanded than in Fig. 25.-27.
Exine sculpture. -28. Polar view of Vigia serrata (Bradem et al. 8376 DAV).-29. Close-up of colpus margin (at
left) and exine sculpture of V serrata (Hoehne 29250 A). Scale bar: = 10 /Am in Figs. 25, 26, 28; = 5 Itm in Figs.
27, 29.
large grains (60-76 jAm E) with an exine ca. 4- of twining woody vines endemic to southeastern
4.5 jim thick, while P. multiglandulosa, P. pen- Brazil.
ninervia (Figs. 23, 24), P. supraglandulosa (Gil- Pollen oblate-spheroidal (P/E = 0.91), 54.5 ,gm
lespie, 1933: figs. 7, 8), and P. verrucosa have P x 60 jm E, tricolpate; amb obtuse-triangular,
medium-sized grains (39-51 jum E) with an exine angulaperturate; colpus broad, margins very un-
ca. 2.5-3 jim. Pollen of both Type 1 and Type 2 even and jagged; exine tectate-perforate, 3-3.5
frequently appear to have small sexinous fragments gm thick, gradually thinning toward the colpus
on the apertural membrane (e.g., Figs. 14, 15, 18, margin; tectum fossulate-foveolate, surface sca-
20, 22). brate.
Romanoa (Figs. 25-27). A monotypic genus Vigia (= Plukenetia) (Figs. 28, 29). A mono-

typic genus of vines and lianas of southeastern large, elliptic in shape with margins uneven, cov-
Brazil, now treated as a species of Plukenetia, P. ered with exine approximately equal in thickness
serrata (Vell.) L. J. Gillespie. to the nonapertural exine, but having a fragmented
Pollen suboblate (P/E = 0.79), 38.5 ,tm P X sexine; exine tectate-perforate, 1-2 ,tm thick, uni-
49 ,tm E, tricolpate; amb obtuse-triangular, an- formly thickened; tectum punctate (i.e>, having
gulaperturate; colpus broad with margins uneven; very small foveolae) and microverrucate; apertural
exine semitectate-reticulate, 2.5-3 tm thick, thin- sexine fragmented with narrow fissures separating
ner at colpus margin; muri often crenate, becoming small, irregularly shaped islands, surface microver-
fragmented and finely gemmate at colpus margin. rucate and sometimes punctate.
Elongate strands of sexine were observed near Although clearly visible under SEM, the aper-
the colpus margin on many grains of both collec- tures are usually not visible under LM (sometimes
tions in SEM (Figs. 28, 29), but were not visible appearing as three less dense, more sculptured
in LM. areas in optical cross section of nonacetolyzed
grains, more frequently visible in acetolyzed grains).
RESULTS SUBTRIBE TRAGIINAE Therefore, even though the grain is aperturate,
shortest and longest axes are given rather than
Acidoton (Figs. 30-35). Five species of
polar and equatorial axes. The apertural sexine
shrubs in the West Indies, Central America, and
appears very similar to the nonapertural sexine,
northern South America. Three species were ex-
but is fragmented into irregularly shaped islands
amined. Two pollen types may be distinguished
separated by usually narrow fissures. Typically, this
based on aperture presence and exine structure.
weaker sexine forms three large, poorly defined
Type 1 (Figs. 30-32). Pollen oblate-spheroi-
colpi (Figs. 36, 39); however, apertural regions of
dal (P/E = 0.88-0.91), 37-40 ,tm P x 42-44
some grains are more irregular and do not form
,tm E, tricolpate; amb subcircular or obscurely
distinct apertures (Fig. 37).
obtuse-triangular; colpus narrow with irregularly
shaped islands of sexine, margin uneven and often Megistostigma (Figs. 40, 41). Five species
indistinct; exine tectate-perforate, ca. 1.5 ,m thick, of twining vines and lianas from southeastern China
uniformly thickened; tectum finely and irregularly to the Philippines and Indonesia. Two species were
foveolate-reticulate, lumina round to slitlike and examined.
usually narrower than the muri, muri often broken Pollen spheroidal to ellipsoid-spheroidal, 48-51
and incomplete, surface microspinulose; apertural ttm S x 52-53 ,tm L, weakly tricolpate, irregularly
sexine islands with surface similar to nonapertural aperturate or sometimes inaperturate; outline cir-
sexine, but more irregular. cular or elliptic; apertural-like areas usually pres-
Acidoton nicaraguensis (synonym: A. vene- ent, either 3(-4), large, elliptic apertures or irreg-
zolanus), the only species in Central and South ularly shaped areas not forming distinct apertures;
America, has pollen of Type 1. exine tectate-perforate, ca. 1 ,tm thick, uniformly
Type 2 (Figs. 33-35). Pollen spheroidal to thickened throughout; tectum punctate and mi-
ellipsoid-spheroidal, 37.5-41.5 ,tm S, 40.5-44 ,tm croverrucate, surface often uneven; sexine of ap-
L, inaperturate; outline circular to broadly elliptic; ertural-like areas fragmented into irregularly shaped
exine tectate-rugulate, ca. 1.5 ,tm thick; rugae islands separated by narrow branched fissures, sur-
short to elongate, sometimes appearing beaded with face microverrucate and sometimes punctate.
slight constrictions at usually regular intervals, sur- The areas of fragmented sexine are less dense,
face smooth, intervening perforations variable in presumably weaker areas of exine that probably
width. function in a manner similar to apertures. In Me-
The two West Indian species examined belong gistostigma malaccense this fragmented sexine
to Type 2. Acidoton microphyllus (Fig. 35) ap- may sometimes be absent or more commonly ran-
pears to have more conspicuously beaded rugae domly distributed, sometimes but not always form-
than A. urens (Figs. 33, 34). ing patterns such as rings (Figs. 40, 41, or similar
to the atypical grain of Cnesmone, Fig. 37). In M.
Cnesmone (Figs. 36-39). A genus of ca. 12
cordata these less dense areas are distributed either
species of twining vines and lianas from south-
in the form of three (or sometimes four), large,
eastern China to the Philippines and Indonesia.
poorly defined colpi as in Cnesmone (Figs. 36, 39)
Four species were examined.
or in a more random pattern.
Pollen spheroidal to ellipsoid-spheroidal, some-
times irregular in shape, 47.5-55 ,tm S, 49.5-56 Pachystylidium (Figs. 42, 43). A monotyp-
,tm L, weakly tricolpate; outline circular or elliptic, ic genus of twining vines distributed from India to
occasionally irregularly obtuse-triangular; aperture the Philippines and Indonesia.

330 Annals of the
~~ ''r*~~~
v"z J
-n
FIGURES3O-35.t Scanning electron micrographs of pollen of Acidoton (subtribe Tragiinae). 30-32. A. nicar-
agulensis (Ortiz 1104 DAy). 30. Equatorial view.- 31. Polar view.-32. Close-up of colpus and exine sculpture;
note presence of sexine islands on the aperture. 33-34. A4. areas. -33. Exine sculpture showing rugae separated
by fossae of variable width.-34. Grain having fossae of mostly narrow width.-35. Grain of A. microphyllus. Scale
bar: = 10 ,um in Figs. 30, 31, 34, 35; = 2 ,um inl Figs. 32, 33.

Pollen oblate-spheroidal (P/E = 0.88-0.92), ing a foveolate-fossulate tectum (Figs. 47, 48) and
30.5-32 Am P X 33-36.5 Am E, weakly triporate; T lessertiana having a reticulate tectum (Figs. 49,
amb subcircular to obtuse-triangular, angulaper- 50).
turate; aperture circular to broadly elliptic, L/W Tragia sect. Ctenomeria (Figs. 51-54).
ca. 1-1.5, covered with exine slightly thinner than Section of two species of southern Africa. Tragia
the nonapertural exine, but with a fragmented sex- capensis examined here.
ine; exine tectate-perforate, 1.5-2 Am thick, uni- Pollen oblate-spheroidal (P/E = 0.89), 31 Am
formly thickened; tectum punctate, microverru- P x 35 Am E, with 3 poorly defined apertures;
cate; apertural sexine fragmented into small amb subcircular; aperture indistinct, appearing as
irregularly shaped islands separated by fissures, an elliptic depressed area covered with a layer of
surface microverrucate and occasionally punctate. exine distinctly thinner than the nonapertural ex-
Pollen grains are similar to those of Cnesmone me, the apertural exine sometimes splitting in an
(Figs. 36-39), but differ in their circular apertures irregular manner following acetolysis; exine tec-
(Fig. 42) that are usually visible under LM as areas tate-perforate, ca. 2 Am thick, much thinner over
of slightly thinner, more sculptured exine. In ad- the aperture; tectum very finely and irregularly
dition, the apertures may sometimes appear as foveolate-reticulate, lumina subcircular and often
depressed areas under SEM (Fig. 42, aperture on narrower than the muri, muri thick, uneven-sur-
right). faced, sometimes broken and incomplete, surface
microspinulose; apertural sexine similar to but more
Platygyna (Figs. 44-46). Seven species of
irregular and finer than the nonapertural sexine,
twining woody vines endemic to Cuba. Three spe-
sometimes irregularly fragmented.
cies were examined.
The foveolate-reticulate tectum is usually con-
Pollen spheroidal or sometimes ellipsoid-sphe-
tinuous over the aperture, with the aperture ap-
roidal, 32-41 Am diam., inaperturate; outline cir-
pearing as an elliptical depression under SEM and
cular to broadly elliptic, often irregularly so; exine
as a much thinner area of exine under LM. The
tectate-perforate, 3-4 Am thick, uniformly thick-
apertures may be considered to be tenuitates, being
ened; tectum reticulate or rugulate with muri or
indistinctly defined areas of thin exine (definition
rugae wider than the usually slitlike intervening
following Erdtman, 1952). Grains may have ap-
perforations, surface smooth.
ertures folded inward in a direction parallel to the
Platygyna hexandra (Fig. 44) and P. leonis
polar axis (Fig. 51). Following acetolysis many
(Fig. 45) have a reticulate tectum with broad muri,
grains had apertures split in an irregular manner
whereas P. parvifolia (Fig. 46) has a rugulate
tectum. (Fig. 54).
Tragia sect. Lassia (Figs. 59, 60). Mono-
Tragia (Figs. 47-74). A genus of ca. 130 typic section of Madagascar consisting of T. scan-
species of herbs, twining vines, and shrubs found dens.
in warm temperate and tropical regions around the Pollen suboblate (P/E = 0.86), 33 P x 38.5
world, particularly abundant in the New World and E, tricolpate; amb subcircular; colpus of narrow to
Africa. Twenty-one species belonging to two sub- medium width often with scattered, irregularly
genera and seven sections were examined. Seven shaped islands of sexine, margin very uneven and
pollen types may be distinguished based on aperture indistinct; exine semitectate-reticulate, ca. 1.5 ,um
condition, aperture morphology and exine struc- thick, uniformly thickened, fragmented at colpus
ture; these types correspond closely with the in- margin; muri microverrucate to obscurely crenate;
frageneric classification system. apertural sexine islands identical in sculpture to
Tragia sect. Bia (Figs. 47-50). Neotropical nonapertural sexine.
section of six species, of which T. lessertiana and Tragia sect. Leptobotrys (Figs. 55-58). The
T. sellowiana were examined here. section consists of two species of the southeastern
Pollen spheroidal or rarely ellipsoid-spheroidal, United States; both species, T urens and T. smallii,
49-54 Am diam., inaperturate; outline circular; were examined.
exine tectate-perforate to semitectate-reticulate, Pollen oblate-spheroidal (P/E = 0.89-0.92),
2.5-3.5 Am thick; tectum foveolate-fossulate or 34-37.5 Am P x 38-40.5 Am E, weakly triporate;
finely reticulate with perforations or lumina often amb subcircular to obtuse-triangular, angulaper-
irregular in size and shape, surface often uneven, turate; aperture circular to very broadly elliptic
microverrucate to densely and irregularly scabrate. with margin indistinct, covered with a fragmented
There is interspecific variation in the size of the exine slightly thinner than the nonapertural exine,
perforations/lumina with Tragia sellowiana hav- often split in an irregular manner; exine tectate-

332 Annalsof the
Ai ~ ~ ~ ~ ~ - i
-~~~~~~~~~~~
V l.A.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~LQ
i~~~ ~ is
FIGURES 36-41. Scanning electron micrographs of pollen of Cnesmone and Megistostigma (subtribeTragiinae).
36-38. Cnesmone anisosepala. -36. Polar view.-37. Atypical irregularlyaperturategrain;note that the fragmented
apertural sexine does not form three apertures.-38. Close-up of aperture margin with fragmented apertural sexine

perforate, 1-1.5 jim thick, uniformly thickened; sometimes be thicker (ca. 2.5 Aim) at mid meso-
tectum punctate, with irregularly shaped raised colpium and thinner toward the aperture margins
areas, surface sparsely to densely microverrucate, (e.g., T. pacifica).
microverrucae variable in size; apertural sexine Tragia sect. Zuckertia (Figs. 67-69). Mono-
consisting of small baculate, rounded, or conical typic section comprising T. bailloniana of Me-
islands, surface smooth or microverrucate. soamerica.
The apertural sexine may be either continuous Pollen oblate-spheroidal (P/E = 0.90), 56 jim
across the aperture (Figs. 55, aperture at top, 57) P x 62.5 jm E, tricolpate; amb obtuse-triangular,
or split in an irregular manner with the underlying sometimes obscurely so, angulaperturate; colpus
membrane protruding through the resulting hole(s) broad with margins uneven and jagged; exine semi-
(Figs. 55, aperture at right, 56, 58). tectate-reticulate, ca. 2,m thick, becoming thinner
Tragia sect. Tagira (Figs. 61-63). Section toward colpus margin; tectum finely reticulate, be-
of ca. 60 species, primarily in dry areas of Africa coming finer near aperture margin, muri scabrate.
but also found in southwest Asia. Four species Section unknown: Tragia novae-hollandiae
examined here, T. adenanthera, T. cord'folia, T. (Figs. 70, 71). A twining vine endemic to Aus-
hispida, and T involucrata. tralia and the only species of Tragia represented
Pollen suboblate to oblate-spheroidal (P/E = there.
0.87-0.88), 31.5-38 jim P X 36-43.5 im E, Pollen oblate-spheroidal to suboblate (P/E =
tricolpate; amb subcircular to obscurely obtuse- 0.88), 36 jim P x 41 jim E, with three poorly
triangular; colpus of narrow to medium width, often defined apertures; amb obtuse-triangular, angulap-
with scattered, irregularly shaped islands of sexine, erturate; aperture circular to broadly elliptic, L/W
margin very uneven and indistinct; exine semitec- ca. 1-1.5, with margin indistinct, covered with a
tate-reticulate, 1.5-3 Aim thick, usually uniformly fragmented exine slightly thinner than the nonaper-
thickened throughout, fragmented at colpus mar- tural exine; exine tectate-perforate, ca. 1.5 jim
gin; muri crenate or microverrucate, often with thick, uniformly thickened; tectum punctate, with
microprojections in 1 or 2 rows; apertural sexine irregularly shaped raised areas, surface microver-
islands identical in sculpture to nonapertural sex- rucate; apertural sexine consisting of small, often
me. conical or baculate islands, surface microverrucate.
Tragia sect. Tragia (Figs. 64-66). A New Tragia subg. Mauroya (Figs. 72-74). Mono-
World section of ca. 55 species of herbs, shrubs, typic subgenus consisting of T ivohibeensis, en-
and twining vines, particularly abundant in dry demic to Madagascar.
subtropical areas. Eight species were examined here: Pollen oblate-spheroidal (P/E = 0.91), 43.5,um
T. chlorocaulon, T mexicana, T. pacifica, T. pel- P x 48,um E, with 3(-4) poorly defined apertures;
tata, T. polyandra, T. ramosa, T. tristis, and T. amb circular to obscurely obtuse-triangular, an-
volubilis. gulaperturate; aperture indistinct, partially covered
Pollen suboblate (P/E = 0.83-0.88), 25-37 with fragments and strands of sexine, margin very
,um P X 30-43.5 ,um E, tricolpate; amb obtuse- indistinct; exine semitectate-reticulate, 1-1.5 jim
triangular, sometimes obscurely so, angulapertur- thick, mostly uniformly thickened; tectum finely
ate; colpus usually broad, often with 1 to several reticulate, muri microverrucate or microspinulose;
widely scattered, irregularly shaped and sized is- apertural sexine fragments and strands similar in
lands of sexine, margin very uneven; exine intec- sculpture to the nonapertural sexine.
tate?-baculate or clavate, 1.5-3 jm thick; baculae The poorly defined apertures appear to be ir-
or clavae usually freestanding, but sometimes close- regular in size and shape, and are often difficult to
ly abutting and appearing coalesced, apex often discern in LM.
flat, with (1-)2-4(-5) microverrucae; apertural
sexine similar in sculpture to the nonapertural sex-
me. Tragiella (Figs. 75-77). Four species of
The exine is generally relatively thin (1.5-2 jm) twining or erect perennial herbs found in southern
and of uniform thickness throughout, but may and eastern Africa. One species examined.
at left and nonapertural sexine at right.-39. Equatorial view of Cnesmone tonkinensis. 40-41. Megistostigma
malaccense (Rahmat si Toroes 1389 A).-40. Irregularly aperturate grain.-41. Close-up showing nonapertural
sexine surrounded by fragmented apertural sexine. Scale bar: = 10 Aimin Figs. 36, 37, 39, 40; = 5 Aimin Figs.
38, 41.

334 Annals of the
i;~~~~~~~~~~~~~~i
JLL
- SX ~~~~~~~~~~~6
FIGURES42-48.3 Scanning electron micrographs of pollen of Pachystylidium, Platygyna, and Tragia sect. Bia
(subtribe Tragiinae). 42-43. Pachystylidium hirsutum (Ramos & Edafio 49201 UC).-42. Equatorialview showing
circular apertures, with aperture at right visible as depressed area.-43. Close-up showing aperture covered with
islands of sexine at bottom center surrounded by nonapertural exine.-44. Grain of Platygyna hexandra. -45.
Exine sculpture of Platygyna leonis. -46. Exine sculpture of Platygyna parvifolia. 47-48. Tragia sect. Bia: T.
sellowiana.-47. Whole grain.-48. Exine sculpture. Scale bar: = 10 ,um in Figs. 42, 44, 47; = 2 ;im in Figs.
43, 45, 46, 48.

w~~
`52
,~~~41t,~~r
FIGURES49-54.3 Scanning electron micrographs of pollen of Tragia sect. Bia and Tragia sect. Ctenomeria.
49-50. Tragia sect. Bia: T. lessertiana. -49. Whole grain.-50. Exine sculpture. 5 1-54. Tragia sect. Ceenomeria:
T: capensis (Kuntze s.n. NY).-51. Equatorial view.-52. Oblique view.-53. Exine sculpture.-54. Equatorial
view of acetolyzed grain with ruptured apertural exine. Scale bar: = 10 ,umin Figs. 49, 51, 52, 54; = 2 Ami in Figs.
50, 53.

336 Annals of the
55 - \kj~~~~~~6
'a?.
2: )~~~~~~~~~O
- **haf W4Ww1_
I to 7, lip-'
A~
* 4 *
FIGURES55-6O.3 Scanning electron micrographs of pollen of Tragia sect. Leptobotrys and Tragia sect. Lassia.
55-58. Tragia sect. Leptobotrys. 55, 56, 58. T. urens. -55. Polar view; note aperture at lower right with a split
apertural sexine. -56. Close-up of aperture with split apertural sexine at lower right and nonaperturalexine at upper
left.-57. Close-up of T. smaliji showing depressed circular aperture.-58. Equatorial view of T. urens showing
circular aperture with ruptured apertural sexine. 59-60. Tragia sect. Lassia: T. scandens. -59. Oblique view. -
60. Close-up of colpus and exine sculpture. Scale bar: = 10 1lm in Figs. 55, 58, 59; = 5 1tm in Figs. 56, 57, 60.

IL4
'v63 I Ai
em~~~~~~~~~~~~~d
63
q(4
as~~
FIGURES61-66. Scanning electron micrographs of pollen of Tragia sect. Tagira and Tragia sect. Tragia. 61-
63. Tragia sect. Tagira: T adenanthera.-61. Oblique view.-62. Equatorial view.-63. Exine sculpture with
uneven and indistinct colpus margin at right. 64-66. Tragia sect. Tragia. -64. Close-up of colpus in polar view of
T. ramosa. -65. Polar view of T. volubilis. -66. Equatorialview of T. volubilis showing colpus with scattered sexine
islands. Scale bar: = 10 tm in Figs. 61, 62, 65, 66; = 5 jsm in Figs. 63, 64.

338 Annals of the
-9 ~~~~~~~~~~~~~~~~V
4 41*i.H4s19.''A 7
1
~ ~ ~~~~~~i
3~~~~~~~~~~~~~~~~~~~~~~i~
4.4.
70 ,.~~~~~~~~~~~~~~~~~~~~~~~~jjjj\
p~~~~~7A
FIGURES67-7 1.3 Scanning electron micrographsof pollen of Tragia sect. Zuckertia and Tragia novae-hollandiae.
67-69. Tragia sect. Zuckertia: T. bailloniana.-67. Polar view.-68. Equatorialview.-69. Exine sculpture. 70-
71. T novae-hollandiae. -70. Polar view. -71. Close-up of circular aperture and exine sculpture. Scale bar: = 10
,gm in Figs. 67, 68, 70; = 5 gm in Figs. 69, 71.
Pollen oblate-spheroidal (P/E = 0.91), 42.5 ,um DISCUSSION

P x 46.5 ,m E, tricolpate; amb subcircular; colpus POLLEN
GENERAL OFTHEPLUKENETIEAE
MORPHOLOGY
narrow, margin very uneven; exine semitectate-
AND PHYLOGENETICIMPLICATIONS
reticulate, ca. 2.5 ,.m thick, mostly uniformly
thickened; muri microverrucate or sometimes ap- A diversity of pollen morphology was found in
pearing crenate with transversely oblong microver- the Plukenetieae, including tricolpate, weakly aper-
rucae. turate, and inaperturate pollen with exine structure
No evidence of apertural sexine islands was seen ranging from tectate, semitectate, to apparently
under LM or in acetolyzed pollen in SEM. intectate (Table 2). Apertures, when present, are

;t ^s
~74}* 94pe* 75)
dh
W ~~~~~~~~ WWVwu
Ar~~~~h
FIGURES 72-77.3 Scanning electron micrographs of pollen of Tragia subgenus Mauroya and Tragiella (subtribe
Tragiinae). 72-74. Tragia subgenus Mauroya: T'. ivohibeensis. -72. Polar view.-73. Equatorial view showing
poorly defined aperture covered with strands and fragments of sexine. -74. Exine sculpture. 75-77. Tragiella
natalensis, acetolyzed grains.-75. Exine sculpture.-76. Polar view.-77. Equatorial view. Scale bar: = 10 jim
in Figs. 72, 73, 76, 77; = 2 jim in Figs. 74, 75.

TABLE 2. Pollen morphology of the Plukenetieae. Aperture condition, tectum morphology (i.e., nonapertural),and presen
each genus, including each section of Tragia and each pollen type (if more than one). The category "absent" also includes
do not form distinct islands on the apertural membrane.
Genus Aperture condition Tectum morphology

Plukenetiinae
Angostyles tricolpate finely foveolate-rugulate absent
Astrococcus tricolpate finely foveolate-rugulate absent-o
Eleutherostigma tricolpate foveolate absent
Haematostemon tricolpate finely foveolate-rugulate absent-o
Plukenetia Type 1 tricolpate foveolate absent
Type 2 tricolpate reticulate absent
Romanoa tricolpate fossulate-foveolate absent
Vigia tricolpate reticulate absent
Tragimae
Acidoton Type 1 tricolpate finely & irregularly scattered
foveolate-reticulate
Type 2 inaperturate rugulate
Cnesmone weakly tricolpate punctate dense co
Megistostigma weakly tricolpate, punctate dense co
irregularly aperturate
or inaperturate
Pachystylidium weakly triporate punctate dense co
Platygyna inaperturate reticulate or rugulate
Tragia sect. Bia inaperturate foveolate-fossulate or finely -
reticulate
T sect. Ctenomeria weakly 3-aperturate finely & irregularly continuo
foveolate-reticulate fragm
T sect. Lassia tricolpate reticulate scattered
T sect. Leptobotrys weakly triporate punctate, uneven-surfaced dense co
T sect. Tagira tricolpate reticulate scattered
T sect. Tragia tricolpate baculate few scat
T sect. Zuckertiana tricolpate finely reticulate absent
T novae-hollandiae weakly triporate punctate, uneven-surfaced dense co
T subg. Mauroya weakly 3-aperturate finely reticulate irregula
Tragiella tricolpate reticulate absent?

simple, i.e., they lack differentiated endoapertures is tricolporate -* tricolpate -- weakly tricolpate,
or ora. An unusual feature is the presence of several triporate or 3-aperturate -+ inaperturate (Fig. 78),
different types of weakly defined apertures. In- the tricolpate condition would still be considered
variant character states include medium to large most primitive in the Plukenetieae even if it is not
size and suboblate to spheroidal shape (i.e., with homologous with the same condition in other Aca-
the polar axis shorter than or equal to the equatorial lyphoideae, but was separately derived from the
axis). tricolporate condition through loss of the endoap-
The majority of Euphorbiaceae have tricolporate erture.
pollen; their compound apertures consist of an out- The genus Dalechampia was treated as a sub-
er colpus and an inner os. Colpate, porate, and tribe of the Plukenetieae by Webster (1994) in his
inaperturate conditions are uncommon in the fam- most recent classification. However, the distinctive
ily outside of the Plukenetieae except within the pollen of Dalechampia (Punt, 1962; Webster &
two subfamilies Oldfieldioideae and Crotonoideae. Webster, 1972: fig. 25) is very different from any
The Oldfieldioideae are characterized by brevicol- pollen type found within the Plukenetieae (as cir-
porate or porate echinate pollen (Levin & Simpson, cumscribed here, i.e., sensu Webster, 1975). Pol-
1994). The Crotonoideae are characterized by the len of Dalechampia is subspheroidal to prolate,
presence of a crotonoid exine structure (Nowicke, tricolporate with thickened equatorial bands of ex-
1994), with the inaperturate condition most com- ine, and coarsely reticulate. The tricolporate ap-
mon and the colpate and porate aperture conditions erture condition of Dalechampia does not support
characteristic of taxa considered by Webster (1994) the hypothesis that the genus was derived from
to be least derived within the subfamily. Both sub- within the Plukenetieae as suggested by Webster
families appear to be distantly related to the Plu- & Webster (1972), given that the independent
kenetieae on the basis of floral, vegetative, and evolution of an endoaperture is a much less likely
pollen exine characters. Inaperturate pollen is not event than its loss. If Dalechampia is instead the
found elsewhere in the Euphorbiaceae and porate sister group of the Plukenetieae, their common
pollen is found rarely and only in distantly related ancestor would presumably have had tricolporate
phyllanthoid taxa (e.g., Phyllanthus L., Hymeno- pollen. This would imply that the tricolpate con-
cardia Wall. ex Lindl.). dition is a synapomorphy defining the Plukenetieae
The Acalyphoideae are characterized primarily and is not homologous with the tricolpate state of
by tricolporate pollen; however, there are also oc- other members of the Acalyphoideae.
currences of tricolpate pollen outside of the Plu-
kenetieae. These include three of five genera in
POLLEN MORPHOLOGYAND PHYLOGENETIC
subtribe Ditaxinae of the Chrozophoreae (Argy-
IMPLICATIONSIN SUBTRIBE PLUKENETIINAE
thamnia P. Browne, Chiropetalum A. Juss., Di-
taxis Vahl) characterized by tricolpate operculate Subtribe Plukenetiinae is relatively uniform in
pollen, subtribe Cephalomappinae of the Epiprineae pollen morphology, characterized by tricolpate pol-
(Cephalomappa Baill.) characterized by brevicol- len with well defined apertures having an uneven,
pate pollen, and tribe Omphaleae (Omphalea) hav- sometimes jagged margin (Table 2). Plukenetia
ing pollen resembling but distinct from Plukenetia (Figs. 14-24) has pollen with broad colpi having
(Punt, 1962; Gillespie, 1988; pers. obs.). Both the very uneven, jagged margins, an amb that is ob-
tricolpate and tricolporate aperture conditions have tuse-triangular and angulaperturate or less often
been described for pollen of Adelia L., Lasiocroton subcircular, and a foveolate or reticulate tectum.
Griseb., Leucocroton Griseb. (all in tribe Adelieae), Two pollen types may be distinguished based on
and Erismanthus Wall. ex Muell. Arg. (tribe Eris- tectum morphology (Table 2); these types corre-
mantheae) (Erdtman, 1952; Punt, 1962; pers. obs.). spond approximately to groups distinguished by
Subtribe Dysopsidinae of the Acalypheae (Dysop- Punt (1962), but are more narrowly defined and
sis Baill.) has pollen with three weakly restricted to Plukenetia (i.e., foveolate Type 1
defined apertures superficially similar to but struc- pollen with Punt's Plukenetia volubilis subtype
turally different from Tragia capensis (Fernindez- and reticulate Type 2 pollen with the Plukenetia
Gonzalez et al., 1994; SuArez-Cervera, pers. verrucosa subtype). Species having Type 1 pollen
comm.). Therefore, based on comparison with the may be subdivided geographically into a neotropical
remainder of the Acalyphoideae the tricolpate con- group characterized by large grains with a thick
dition would appear to be most primitive in the exine that tapers toward the margin and a paleo-
Plukenetieae. Given that the most plausible direc- tropical group characterized by medium-sized grains
tion of evolution of aperture condition in the tribe having a thinner exine that is uniformly thick or

342 Annals of the
J ( h
FIGURE 78.3 Evolution of aperture condition in the Plukenetieae: a hypothetical character state tree. Note that
only one aperture is shown in equatorial view, and that only apertural condition is schematically illustrated,not details
such as aperture margin morphology.-a. Ancestral tricolporate condition (Dalechampia, most Acalyphoideae).-b.
Tricolpate (all Plukenetiinae, Tragia sect. Zuckertia).-c. Tricolpate with scattered sexine islands on apertural
membrane (Tragia sect. Tragia, Tragia sect. Tagira, Acidoton Type 1). -d. Weakly tricolpate, apertures with dense
covering of small sexine islands (Cnesmone, Megistostigma).-e. Irregularlyaperturate, aperturalareas with a dense
covering of small sexine islands (Megistostigma).-f. Inaperturate(Megistostigma).-g. Weakly triporate, apertures
with dense covering of small sexine islands (Pachystylidium, Tragia sect. Leptobotrys, Tragia novae-hollandieae). -
h. Inaperturate (Acidoton Type 2, Platygyna, Tragia sect. Bia).-i. Weakly 3-aperturate, apertural areas covered
with thin layer of exine, usually visible as depressions (Tragia sect. Ctenomeria).-j. Weakly 3-aperturate, apertural
areas covered with strands of sexine (Tragia subgenus Mauroya).
thicker at the aperture margin. Species having be distinguished by its trilocular ovary and 5-parted
Type 2 pollen, known only from the Neotropics, pistillate calyx, both plesiomorphic characters in
may also be subdivided into two groups using the the subtribe. Romanoa tamnoides appears to be
above criteria. The two pollen types and subgroups either the sister taxon of Plukenetia, which is char-
appear to have a phylogenetic basis and reflect acterized by a 4-locular ovary and 4-parted calyx,
current ideas of species relationships in the genus or its most plesiomorphic member.
(Gillespie, 1993). Among neotropical species, pol- Angostyles (Figs. 1-3), Astrococcus (Figs. 4-
len morphology, together with degree of style fusion 6), and Haematostemon (Figs. 9-13) share a very
and androecium morphology, may be used to define similar pollen morphology, which is distinct in col-
major species groups. pus and tectum morphology from that of the other
Pollen morphology of the monotypic genera genera of subtribe Plukenetiinae described above
Eleutherostigma (Figs. 7, 8) and Vigia (Figs. 28, (Table 2). Pollen is subcircular and not distinctly
29) fits well within the range of variation found in triangular, with a very finely rugulate, microver-
Plukenetia, thus strongly supporting their recent rucate tectum. Colpi are narrower with an uneven
synonymy under Plukenetia (Gillespie, 1993). but not distinctly jagged margin. The three genera
Eleutherostigma has pollen of Type 1 and appears share a trilocular ovary and a tree or shrub habit,
to be very closely related to the other neotropical very different from the vine or liana habit of PMu-
species included in Type 1, while Vigia has pollen kenetia and Romanoa. Both Astrococcus and
of Type 2 that is medium sized, similar to P. Haematostemon have unusually thickened aper-
verrucosa, P. penninervia, and related species. ture margins with the upper and lower exine layers
Romanoa (Figs. 25-27) also has pollen very separating to form an elongate chamber (similar to
similar to that of Plukenetia, but differs in its a vestibulum). Pollen characters together with an
fossulate-foveolate tectum. Although morphologi- androecium of four stamens, a unique character
cally very similar to Plukenetia, the genus may in the subtribe, suggest their very close relation-

ship. In contrast, Angostyles lacks the thickened distinct pollen types, differing in aperture presence
aperture margin and elongate chamber and bears and exine sculpture. The tricolpate grains of the
staminate flowers with numerous stamens. Thus mainland neotropical species, A. nicaraguensis,
while Angostyles, Astrococcus, and Haematoste- have colpi with uneven margins and apertural sex-
mon form a distinct group, the latter two are most ine islands (Figs. 30-32). The inaperturate pollen
closely related and Angostyles is the least derived of the West Indian species (Figs. 33-35) appears
member sharing with Plukenetia the plesiomorphic more similar to pollen of its geographical neighbor
characters of numerous stamens and absence of Platygyna (Figs. 44-46), particularly to P. par-
an elongate chamber. Punt (1962) also remarked vifolia, than to its mainland congener. Interest-
that pollen morphology of Angostyles is interme- ingly, P. parvifolia appears morphologically in-
diate, sharing with Astrococcus and Haematoste- termediate between the two genera, having an
mon a "tectum perforatum" (as opposed to a "psi- intermediate stamen number and a staminate re-
late" tectum as described for other species in the ceptacle that is glabrous like Acidoton but globose
Plukenetia volubilis subtype), but apparently lack- like Platygyna. This suggests that the mainland
ing a "margo" (defined as a prominent or depressed species is not the sister taxon of the West Indian
margin) as in Plukenetia (except P. conophora). species of Acidoton, and therefore should be treat-
ed as the distinct genus Gitara Pax & K. Hoffm.
(as originally considered by Pax & Hoffmann,
POLLEN MORPHOLOGYAND PHYLOGENETIC
1924). An alternative hypothesis would be the or-
IMPLICATIONSIN SUBTRIBE TRAGIINAE
igin of the inaperturate condition from the tricol-
Subtribe Tragiinae exhibits much greater diver- pate condition within Acidoton, and independently
sity in pollen morphology than subtribe Plukene- from the inaperturate condition in Platygyna;
tiinae. Pollen is tricolpate, inaperturate, or with however, given their very similar pollen including
poorly defined apertures. Exine sculpture includes tectum morphology, this hypothesis would seem
punctate, foveolate, rugulate, reticulate, and bac- less probable.
ulate conditions (Table 2). Pachystylidiium and The three Indomalaysiangenera of subtribeTra-
Platygyna are characterized by unique pollen types; giinae, Cnesmone (Figs. 36-39), Megistostigma
Acidoton includes two distinct pollen types. The (Figs. 40, 41), and Pachystylidium (Figs. 42, 43),
large genus Tragia includes seven very distinct form a distinct group based on pollen morphology.
pollen types, each characteristic of one or more They are characterized by weakly defined aper-
sections with each section having a uniform pollen tures or apertural regions (sometimes absent in
morphology. Megistostigma), a tectate-punctate exine with su-
Pollen of the West Indian genus Platygyna pratectal microverrucae, and an apertural sexine
(Figs. 44-46) is inaperturate with a reticulate or that is fragmented into small islands (Table 2). The
rugulate tectum. No evidence was seen of small primary differences among the genera are in the
circular endexine thickenings observed by Punt shape and size of the apertural sexine regions.
(1962) on acetolyzed grains of P. hexandra. The Pachystylidium (Figs. 42, 43) has three circular,
generic status of Platygyna has been questioned porelike apertures, whereas Cnesmone (Figs. 36,
(Liogier, 1971; Borhidi et al., 1973); only two floral 39) typically has three elliptic colpuslike apertures.
characters, a globose or convex staminate recep- In Megistostigma (Fig. 40), the apertural condi-
tacle and thickened papillose styles, separate the tion varies from three (or four) colpuslike apertures
genus from Tragia. Palynologically, Platygyna to more randomly distributed apertural regions or
appears to be quite distinct from Tragia with the sometimes inaperturate. Megistostigma is char-
exception of the neotropical section Bia (Figs. 47- acterized by having an apertural condition that
50), the only section characterized by inaperturate varies both between and within species (including
pollen. Platygyna differs from section Bia in its within a single sample); M. cordata has pollen with
smaller pollen grains with a coarser exine sculpture. either three colpuslike apertures as in Cnesmone
Differences in inflorescence architecture, presence or with randomly distributed irregular apertural
of disc segments or a globose receptacle in the regions, while M. malaccense is weakly and irreg-
staminate flowers and style morphology between ularly aperturate or inaperturate. Pollen evidence
these two taxa would seem to preclude a close supports the hypothesis based on floralmorphology
relationship, suggesting that the similarity in pollen that Megistostigma and Cnesmone are sister taxa.
morphology, particularly the inaperturate condi- In fact, the two genera may not be as distinct as
tion, is due to convergence. previously thought; the traditional distinctions of
The genus Acidoton was found to contain two style morphology and presence of a staminate ap-

344 Annals of the
pendage appear to break down as more species are areas partially covered with fragments and strands
recognized and described. Pollen characters, as of sexine (and usually not visible under LM). The
described above, cannot be used to separate the tectum, both apertural and nonapertural, is retic-
two genera. Since pollen of Pachystylidium is in- ulate but finer than in Tragiella and Tragia sect.
termediate between pollen of Cnesmone and Me- Tagira. The species appears most similar in veg-
gistostigma and pollen of several species of Tragia etative and floral morphology to the endemic Mad-
(T. novae-hollandiae and section Leptobotrys), agascan section Agirta, differing primarily in style
pollen morphology does not strongly support either morphology. Pollen of Tragia sect. Agirta needs
recognition of Pachystylidium as a distinct mono- to be examined to confirm this suggested relation-
typic genus (following Pax & Hoffmann, 1919; ship.
Airy Shaw, 1969, 1975) or as a species of Tragia The second pollen type unique among African
(following Webster, 1975). Pollen evidence does Tragiinae is found in the southern African taxon
suggest a relationship with its geographical neigh- Tragia sect. Ctenomeria (Harv.) Benth. While re-
bors, Megistostigma and Cnesmone, despite a very sembling the majority of African Tragiinae in hav-
different staminate flower morphology. ing pinnatifid pistillate sepals (a feature found only
The Old World, predominantly African, taxa, in the Old World), the section has a very distinct
Tragia sect. Tagira Muell. Arg. (Figs. 61-63), androecium of numerous stamens (30-50) with
Tragia sect. Lassia (Figs. 59, 60), and Tragiella highly elongate anthers. Palynologically very dis-
(Figs. 75-77) were found to share a similar pollen tinct also, section Ctenomeria is characterized by
morphology (Table 2). Pollen is tricolpate with a weakly defined apertures (tenuitates), which may
reticulate tectum, a very uneven aperture margin, split in an irregular manner, and a reticulate tec-
and scattered islands of sexine often present on the tum, which is often continuous but much thinner
apertural membrane (apparently absent in Tra- across the aperture (Figs. 51-54). This pollen type
giella). Pollen evidence suggests that the three is unique in the tribe and cannot be easily related
taxa are closely related and supports Webster's to other types, emphasizing the distinctness of sec-
(1975) treatment of Tragiella as a synonym of tion Ctenomeria. A somewhat similar pollen type
Tragia (a new section within Tragia would be is found in the isolated, monotypic South American
necessary to accommodate the species of Tragiel- genus Dysopsis (tribe Acalypheae) (FernAndez-
la), rather than as a distinct genus (following Rad- Gonzalez et al., 1994); however, similarity with
cliffe-Smith, 1982, 1987). Section Lassia is mor- Tragia sect. Ctenomeria is most likely due to con-
phologically very similar to Tagira and may not vergence.
be distinct from that section; Lassia is distinguished Four distinct pollen types are found among neo-
by a single apomorphic androecial character. tropical species of Tragia. Section Bia is the only
Sphaerostylis (not examined here) also has tri- section of Tragia characterized by inaperturate
colpate pollen with a reticulate tectum similar to pollen (Figs. 47-50). Pollen morphology together
the above taxa according to observations by Punt with inflorescence architecture and staminate flow-
(1962). This evidence is consistent with Croizat's ers having disc segments and five to many stamens
(1941) hypothesis of a relationship with Tragiella (5-20) emphasize the distinctness of section Bia,
(though differences in calyx and foliar morphology, as was also pointed out by Punt (1962). A second
as pointed out by Radcliffe-Smith (1987), would pollen type is represented by Tragia bailloniana
provide evidence against combining the two genera) (Figs. 67-69) belonging to the monotypic section
rather than with Megistostigma (following Pax & Zuckertia (Baill.) Muell. Arg. The tricolpate pollen
Hoffmann, 1919). of T. bailloniana is remarkably similar to pollen
Two African taxa of Tragia were found to have of Plukenetia (Figs. 14-24) and Romanoa (Figs.
unique pollen types very different from the above 25-27), differing from Plukenetia Type 2 pollen
African Tragiinae. Tragia ivohibeensis is an en- primarily in its more finely reticulate tectum and
demic Madagascan species for which LUandri (1971) muri that are often scabrate but not crenate, and
created the monotypic subgenus Mauroya, appar- from Romanoa only in the larger size of the per-
ently because he thought it intermediate between forations. Baillon (1858) originally described the
several genera including Tragiella and Sphaero- species as Zuckertia cordata and pointed to a
stylis, and two sections of Tragia, Agirta, and similarity with Romanoa, while Miller & Webster
Ratiga. Pollen morphology of T ivohibeensis (Figs. (1967) suggested that the species was one of the
72-74), however, does not resemble either Tra- most primitive in Tragia. Since presence of stinging
giella or any species of Tragia examined. The hairs and a slender 3-branched style places T
apertures (usually three) are very poorly defined bailloniana in Tragia and not in subtribe Pluke-

netiinae, pollen evidence supports the hypothesis morphology, and apertures densely covered with
that the species is one of the least derived members sexine islands support Miller & Webster's (1967)
of Tragia. The section Zuckertia pollen type ap- conclusions based on staminate flower morphology
pears to represent the plesiomorphic condition in and LM observations of pollen that Mueller's sec-
subtribe Tragiinae based on outgroup comparison tion Leptobotrys is valid and should not be included
with subtribe Plukenetiinae. in section Tragia (as done by Pax & Hoffmann,
The majority of New World species of Tragia 1919).
belong to section Tragia, which appears to be Curiously, pollen of Tragia sect. Leptobotrys is
characterized by a single unique pollen type. Pollen most similar to pollen of T. novae-hollandiae (Figs.
of all species examined is tricolpate with an un- 70, 71), the only species of Tragia known from
usual, apparently intectate sexine consisting of bac- Australia (section undetermined; although Pax &
ulate or clavate sculptural elements (Figs. 64-66). Hoffmann (1919) included the species in section
Pollen is similar to Tragia sect. Tagira in having Leucandra, it is anomalous both in that section
apertures with very uneven margins and scattered and section Tragia). The two taxa share a very
islands of sexine on the apertural membrane, but similar aperture condition, tectum morphology, and
differs in exine structure and broader colpi with obtuse-triangular shape (Table 2). Together they
usually fewer sexine islands. While described as most closely resemble the southeast Asian genus,
operculae by Miller & Webster (1967) on the basis Pachystylidium, differing primarily in details of
of LM observations, these apertural sexine islands tectum morphology, both apertural and nonaper-
are irregular in shape, size, and position, and there- tural. Airy Shaw (1969) pointed to a relationship
fore cannot be considered as operculae (according between T novae-hollandiae and Pachystylidium
to Erdtman's (1952) definition of an operculum as based on the shared state of subsessile anthers and
a "thickening of measurable bulk and clearly de- suggested that the species is transitional between
fined of an aperture membrane (circular in pori, Pachystylidium and Tragia. In addition, Tragia
elongate in colpi . . .)"). Although only eight species sect. Leptobotrys and Pachystylidium share a
of this large section were examined, LM obser- stamen number of two, an unusual condition in the
vations by Punt (1962), Miller & Webster (1967), tribe. Whether similarity in pollen morphology be-
and R. Urtecho (pers. comm.) suggest that this tween these three disjunct taxa is due to homology
pollen type is characteristic of section Tragia and or convergence needs to be examined further.
represents the only occurrence of intectate pollen
(sensu Walker & Doyle, 1975, i.e., excluding the
APERTURE EVOLUTIONIN TRIBE PLUKENETIEAE
semitectate condition considered by Punt, 1962,
as intectate) in the tribe Plukenetieae. Pollen synapomorphies defining the Plukene-
Several sections included here in section Tragia tieae are tricolpate aperture condition and uneven
are sometimes treated as distinct sections (Pax & aperture margins. Tricolpate aperture condition is
Hoffmann, 1919, 1931). These include section primitive in the tribe based on outgroup comparison
Ratiga Muell. Arg. (which includes Tragia chlo- with Dalechampia and the remainder of the sub-
rocaulon, T. mexicana, and T. tristis), section family Acalyphoideae. Throughout the Plukene-
Leucandra (T polyandra, T ramosa), and section tieae aperture margins are uneven and often appear
Leptorhachis (Klotzsch) Muell. Arg. (resurrected fragmented, jagged, or frayed. The character state
by Milgura de Romero & Gutierrez de Sanguinetti tree in Figure 78 illustrates one hypothesis of the
(1989) for several South American species, in- evolution of aperture condition in the Plukenetieae,
cluding T polyandra, but treated by Pax and and will be discussed in greater detail below.
Hoffmann under section Leucandra). Since these In subtribe Plukenetiinae aperture condition is
sections share the same pollen type as section Tra- uniformly tricolpate (Table 2, Fig. 78b). Variation
gia (sensu Pax & Hoffmann; including T. pacifica, in aperture morphology is primarily in colpus size
T. peltata, and T. volubilis), pollen morphology and shape, presence of an elongate chamber within
does not support their recognition as distinct sec- the exine next to the colpus, and presence of an
tions. apertural sexinous membrane.
The fourth pollen type among New World Tra- In subtribe Tragiinae there has been an unusual
gia is found in Tragia sect. Leptobotrys (Figs. radiation in aperture condition and morphology
55-58). Pollen has three poorly defined circular (Table 2). There is a distinct trend toward less well
apertures with a very indistinct margin and covered defined apertures and ultimately toward loss of
with small, often conical or baculate islands of apertures in several evolutionary lines. Aperture
sexine. The distinct apertural condition, tectum margins have become more irregular and less de-

346 Annals of the
fined, and fragments or distinct islands of sexine CONCLUSIONS

are often present on the apertural membrane.
The majority of species of Tragia are tricolpate; The use of SEM in the present study of pollen
all of these species, with the exception of T bail- morphology of the Plukenetieae has provided a
loniana, have scattered islands of sexine on the much greater number of systematically useful char-
aperturalmembrane and very uneven margins (Fig. acters than was possible with LM alone. In partic-
78c). A single species of Acidoton (Type 1) also ular, details of tectum morphology were much more
has pollen of this type. highly resolved and enabled detection of unusual
Three different types of weakly defined aper- apertural conditions and morphology. Several taxa,
tures are found in the Tragiinae, each presumably not considered to represent distinct pollen types in
originating independently from the tricolpate con- Punt's (1962) study based on LM alone, were
dition. One type consists of apertural areas densely distinct when viewed under SEM. These include
covered with numerous small islands of sexine and Tragia capensis grouped by Punt with the PMu-
with an exine equal in thickness or only slightly kenetia volubilis subtype, Tragiella and Tragia
thinner than the nonapertural exine. These areas sect. Tagira grouped with the Plukenetia verru-
may be either elliptic (Fig. 78d; Cnesmone, Fig. cosa subtype, and Acidoton included in the Cnes-
39, and Megistostigma), circular (Fig. 78g; mosa type with Cnesmone and Megistostigma.
Pachystylidium, Fig. 42, Tragia sect. Leptobo- The tricolporate, coarsely reticulate pollen of
trys, Figs. 57, 58, and T. novae-hollandiae, Fig. Dalechampia supports the hypothesis of Dale-
71), or irregular in shape (Fig. 78e; Megistostig- champia as the sister taxon of the Plukenetieae
ma, Fig. 40). A second type of weakly defined (as circumscribed here), rather than being derived
aperture is represented by Tragia sect. Cteno- from within the tribe. Webster's (1994) treatment
meria (Fig. 51), which has apertures covered with of the genus as a subtribe within the Plukenetieae
a continuous or sometimes fragmented, distinctly is consistent with either hypothesis.
thinner exine, often visible as depressed areas (Fig. Pollen evidence is consistent with Webster's
78i). Tragia subg. Mauroya (Fig. 73) represents (1975) division of tribe Plukenetieae into two sub-
a third type characterized by very weakly defined tribes, Plukenetiinae and Tragiinae, but does not
apertures covered with strands of sexine continuous support Pax & Hoffmann's (1919) division into
with and identical in sculpture to the nonapertural four informal groups. Of these groups only As-
exine (Fig. 78j). trocciformes (comprising Astrococcus and Hae-
Inaperturate pollen appears to have evolved at matostemon) is monophyletic, Tragiiformes (Tra-
least twice in subtribe Tragiinae. In the Old World, gia) is paraphyletic, while Plukenetiiformes (most
inaperturate pollen (Fig. 78f) evolved via weakly Plukenetiinae plus Acidoton and Platygyna) and
defined colpate (Fig. 78d) and irregularly apertur- Sphaerostyliformes (all other Tragiinae) appear to
ate grains (Fig. 78e) and is found only in Megis- be polyphyletic.
tostigma (note that apertural condition is variable Pollen morphology supports a subdivision of sub-
within the genus and within species, e.g., M. ma- tribe Plukenetiinae into an arborescent group (An-
laccense has both irregularly weakly aperturate gostyles, Astrococcus, and Haematostemon) and
and inaperturate grains, whereas M. cordata has a lianous group (Plukenetia and Romanoa). Also
both weakly tricolpate and irregularly aperturate supported is the recent synonymy of Vigia and
grains). In the New World, inaperturatepollen (Fig. Eleutherostigma under Plukenetia (Gillespie,
78h) originated one or more times most probably 1993), and the relationship of Astrococcus and
from tricolpate pollen having scattered sexine is- Haematostemon as sister taxa.
lands on the apertural membrane (Fig. 78c) and Subtribe Tragiinae exhibits an exceptionally di-
is found in three taxa, Platygyna (Fig. 44), Acido- verse pollen morphology, with much of the vari-
ton Type 2 (Figs. 34, 35), and Tragia sect. Bia ation present in the large genus Tragia. Pollen
(Figs. 47, 49). An alternative hypothesis would be morphology, for the most part, supports the sec-
origin directly from tricolpate grains lacking ap- tional classification of Tragia. The two largest sec-
ertural sexine islands (Fig. 78b). Acidoton Type 2 tions, neotropical Tragia and paleotropical Tagira,
(and possibly Platygyna) pollen most likely orig- have tricolpate pollen with scattered sexine islands
inated from Acidoton Type 1, i.e., pollen having on the apertural membrane, but are easily distin-
narrow colpi with numerous sexine islands, whereas guished based on exine morphology. Of the re-
there is no strong evidence either way for the origin maining taxa, sections Bia, Ctenomeria, Lepto-
of inaperturate pollen in Tragia sect. Bia. botrys, and Zuckertia, and subgenus Mauroya are

each characterized by a unique pollen Itype, pro- hypotheses of phylogenetic relationships suggested
viding evidence that they are indeed distinct natural in this paper need to be further tested by means
groups. Pollen evidence does not support Lassia of cladistic studies using floral, vegetative, and pol-
as a section distinct from Tagira, but does support len characters. Taxa sharing a similar pollen mor-
the inclusion of sections Leucandra, Leptorhachis, phology, but not previously thought to be closely
and Ratiga within section Tragia. The hypothesis related, need to be more thoroughly examined to
of section Zuckertia as a plesiomorphic member determine if these similarities are homologous or
of Tragia is supported. the result of convergence. A more complete un-
The remaining genera in subtribe Tragiinae, for derstanding of the phylogeny of subtribe Tragiinae
the most part, are characterized by unique pollen is recommended prior to making specific changes
types distinct from Tragia. The southeast Asian in the generic and sectional classification.
taxa, Megistostigma and Cnesmone, form a dis-
tinct group based on pollen morphology that is most
LITERATURE
CITED
similar to pollen of Pachystylidium. Pollen evi-
dence is consistent with the status of Platygyna AIRY SHAW, H. K. 1968. Notes on Malesian and other
Asiatic Euphorbiaceae. XCII. Note on the affinity of
as a genus distinct from Tragia. Acidoton includes Ramelia Baill. Kew Bull. 21: 401-403.
two different pollen types, inaperturate and tricol- . 1969. Notes on Malesian and other Asiatic
pate; the close similarity of inaperturate Type 2 Euphorbiaceae. CXII. Notes on the subtribe Pluke-
pollen to pollen of Platygyna suggests that Acid- netiinae Pax. Kew Bull. 23: 114-121.
. 1974. Notes on Malesian and other Asiatic
oton may not be monophyletic. Tragiella has pol-
Euphorbiaceae. CXII. New or noteworthy species of
len very similar to Tragia sect. Tagira, and would Bocquillonia Baill., with a revised key to the genus.
perhaps be best considered a section of Tragia Kew Bull. 29: 315-322.
related to section Tagira. . 1975. The Euphorbiaceae of Borneo. Kew
Pollen and floral morphological evidence indi- Bull. Add. Ser. IV.
ARMBRUSTER,W. S. 1994. Early evolution of Dale-
cate that a major reorganization of generic delim- champia (Euphorbiaceae):Insights from phylogeny,
itations may be necessary in the Tragiinae to better biogeography, and comparative ecology. Ann. Mis-
reflect phylogenetic relationships. Tragia appears souri Bot. Gard. 81: 302-316.
to be a highly paraphyletic genus as presently BAILLON,H. E. 1858. Etude generale du groupe des
Euphorbiacees. Victor Masson, Paris.
circumscribed. Most of the remaining Tragiinae
BENTHAM,G. 1880. Euphorbiaceae. In: G. Bentham &
genera appear to be derived with respect to Tragia J. D. Hooker (editors), Genera Plantarum ad Ex-
and defined on the basis of unusual and presumably emplaria Imprimis in Herbariis Kewensibus Servata
apomorphic style and androecium characters (and Definita 3: 239-340. Lovell Reeve, London.
also by apomorphic pollen characters). Pollen ev- BORHIDI,A., J. ACUNA& G. MUNIZ. 1973. New plants
in Cuba III. Acta Bot. Acad. Sci. Hung. 19: 37-45.
idence frequently suggests a close relationship CROIZAT,L. 1941. The tribe Plukenetiinae of the Eu-
among geographical neighbors, a relationship that phorbiaceae in eastern tropical Asia. J. ArnoldArbor.
is not reflected in the current classification. To 22: 417-431.
ensure a phylogenetic classification, it may be nec- ERDTMAN,G. 1952. Pollen Morphologyand Plant Tax-
onomy. Angiosperms. Almqvist & Wiksell, Stock-
essary to subdivide Tragia and recognize certain
holm.
sections, such as Bia and Ctenomeria, as distinct 1966. Pollen Morphology and Plant Taxon-
genera (as they have been treated in the past, e.g., omy. Angiosperms (corrected reprint of the 1952
Baillon, 1858), since these sections are as distinct edition with a new addendum). Haffner, New York.
as most Tragiinae genera. An alternative but more FERNANDEZ-GONZALEZ, D., M. SUAREZ-CERVERA & A. LE
THOMAS. 1994. Ultrastructure du pollen de Dy-
cumbersome approach would be to combine most sopsis Baill.: Relations phylogenetiques au sein des
Tragiinae genera into Tragia with a complex sys- Acalypheae (Euphorbiaceae). Acta Botanica Gallica
tem of subgenera and sections to indicate relation- 141 (in press).
ships. The level at which taxa such as Tragiella, GILLESPIE,L. J. 1988. A revision and phylogenetic
and Platygyna are recognized analysis of Omphalea (Euphorbiaceae).Unpublished
Pachystylidium,
Ph.D. Dissertation. University of California, Davis.
will depend on which approach is followed. 1993. A synopsis of neotropical Plukenetia
The present study should not be regarded as (Euphorbiaceae)includingtwo new species. Syst. Bot.
exhaustive; some taxa need to be more thoroughly 18: 575-592.
surveyed, particularly the two largest sections of GREUTER,W. ET AL. (editors). 1988. InternationalCode
of Botanical Nomenclature. Regnum Veg. 118.
Tragia, Tagira and Tragia (to verify that only a KOHLER, E. 1965. Die Pollenmorphologie der biovu-
single pollen type is found in each), while several laten Euphorbiaceaeund ihre Bedeutung fur die Tax-
Madagascan taxa still need to be examined. The onomie. Grana Palynol. 6: 26-120.

348 Annals of the
LEANDRI, J. 1971. Un sous-genre malgache nouveau zenfamilien, ed. 2, 19c: 11-233. W. Engelmann,
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deae (Euphorbiaceae).Ann. Missouri Bot. Gard. 81: RADCLIFFE-SMITH, A. 1980. A note on Romanoa (Eu-
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MILLER, K. I. & G. L. WEBSTER. 1967. A preliminary BELL. 1974. Vascular Plant Systematics. Harper
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Pollen Morphology and Phylogeny of the Tribe Plukenetieae (Euphorbiaceae)

Author(s): Lynn J. Gillespie

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ANN. MISSOURIBOT. GARD. 81: 317-348. 1994.

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Species Collection Location

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Species Collection Location

Acidoton nicaraguensis (Hemsley) Webster Ortiz 1104, DAV Nicaragua

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Species Collection Location

T capensis Thunb. Ecklon & Zeyher s.n., US 1170932 South Africa

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3 Voucher informationis in Table 1 unless given in the caption.

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This content downloaded from 139.184.14.159 on Sat, 7 Feb 2015 11:49:57 AM

This content downloaded from 139.184.14.159 on Sat, 7 Feb 2015 11:49:57 AM

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Pollen oblate-spheroidal (P/E = 0.91), 42.5 ,um DISCUSSION

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Genus Aperture condition Tectum morphology

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fined, and fragments or distinct islands of sexine CONCLUSIONS

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