A New Section and Two New Species of Tragia (Euphorbiaceae) from
the Venezuelan Guayana and French Guiana
Lynn J. Gillespie
Botany Department NHB 166, Smithsonian Institution, Washington, D.C. 20560, U.S.A.
Present address: Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D,
Ottawa, Ontario KIP 6P4, Canada
ABSTRACT. A new section and two new species of
Tragia are described and illustrated. Tragia guay-
anensis from Amazonas, Venezuela, is unique in the
genus in having filaments entirely fused into an
elongate staminal column and 5-colpate pollen. The
new section Monadelphae is described to accom-
modate this unusual species. Tragia tabulaemon-
tana from French Guiana belongs to section Tragia
and is characterized by oblanceolate or narrowly
elliptic leaf blades that are narrowly cordate at the
base, stamens that are highly dilated at their base,
and long-stipitate glandular trichomes on petioles,
inflorescence axes, and ovaries. A key to the five
sections of Tragia in the New World and a key to
the five species of Tragia in the Guayana Region
are given.
Tragia L. is the largest genus of tribe Plukene-
tieae (Euphorbiaceae: subfamily Acalyphoideae) with
approximately 130 species. This genus of twining,
scandent, and erect herbs and subshrubs is widely
distributed in tropical to warm temperate areas in
the New World, Africa, southern Asia, and Austra-
lia, but is most abundant in drier areas of Africa
and the New World. Tragia belongs to subtribe
Tragiinae, which is distinguished from subtribe Plu-
kenetiinae by its urticating trichomes, trilocular ova-
ry, and absence of foliar glands.
Two new species of Tragia were encountered
while preparing treatments of tribe Plukenetieae for
the Guianas (Gillespie & Armbruster, in prep.) and
for the Flora of the Venezuelan Guayana (Gillespie,
in prep.). A total of five species of Tragia are now
known from the Guayana Region (an area encom-
passing the Guianas, the Venezuelan Guayana, and
adjacent areas of Amazonian Brazil north of the
Amazon River; Mori, 1991; Gillespie, 1993). One
of these new species, T. guayanensis, is so distinct
that it does not fit into any of the currently rec-
ognized sections. A new section is described to ac-
commodate this unusual species and is designated
NovoN 4: 330-338. 1994.
Monadelphae, alluding to its unique androecium
within Tragia.
Nine sections of Tragia are currently recognized
(Gillespie, 1994; see also Pax & Hoffmann, 1919).
These sections are for the most part morphologically
very distinct from one another and each is char-
acterized by a unique pollen type (with the exception
of sections Tagira and Lassia, which share a pollen
type) (Gillespie, 1994). Among the five New World
sections, section Tragia is the largest and most
widespread with 40-50 species distributed from the
southern United States to Argentina. The remaining
sections are much smaller and geographically more
restricted; in addition to Monadelphae, these in-
clude Bia (Klotzsch) Mueller Argoviensis (ca. 6 spe-
cies from Panama to Argentina), Leptobotrys
(Klotzsch) Mueller Argoviensis (2 species in the
southeastern United States), and Zuckertia (Baillon)
Mueller Argoviensis (1 species in Mesoamerica).
Among the four Old World sections Tagira Mueller
Argoviensis is the largest with 40-50 species wide-
spread in Africa and southern Asia. The remaining
sections include Ctenomeria Harvey (2 species in
southern Africa), Agirta Baillon (4 species in Mad-
agascar), and Lassia Baillon (monotypic in Mada-
gascar) [also in Madagascar is the single described
monotypic subgenus Mauroya, which is probably
best recognized at the sectional level].
Tragia as presently circumscribed is distin-
guished from other members of subtribe Tragiinae
by a combination of plesiomorphic character states,
such as slender, only partly connate styles and a
flat, glabrous staminate receptacle. The other Tra-
giinae genera are mostly defined by one or more
character states that appear to be derived with re-
spect to Tragia. These states include massive free
styles (distinguishing Cnesmone), styles entirely con-
nate into a massive column (Megistostigma and
Sphaerostylis), convex hairy receptacle (Platy-
gyna), and anther connective with apical tuft of
stinging hairs (Acidoton).


Volume 4, Number 4
1994
The circumscription and number of both sections
of Tragia and genera of Tragiinae is open to question
and needs further study. A major reorganization
within subtribe Tragiinae may be necessary to better
reflect phylogenetic relationships (Gillespie, 1994).
This may include recognition of several sections of
Tragia, such as Bia, Ctenomeria, and Monadel-
phae, at the generic level.
KEY TO THE NEOTROPICAL SECTIONS OF TRAGIA
1. Filaments completely connate into a central
elongate column bearing a cluster of sessile an-
thers; pollen 5-colpate, staminate sepals 5; in-
florescences unisexual, racemose ...........
.... ................ .. section M onadelphae
1. Filaments free or rarely partly connate; pollen
tricolpate, weakly triporate, or inaperturate; sta-
minate sepals 3-5(-6); inflorescences bisexual,
racemose or branched ...................... 2
2. Inflorescence consisting of a racemose stami-
nate main axis and a single elongate basal branch
bearing 5-20 pistillate flowers; stamens 8-40+;
staminate disc segmented or sometimes absent
... .................................... 3
2. Inflorescence racemose, with 1(-2) pistillate
flower(s) at the basal 1(-9) node(s); stamens
(1-)2-5(-22); staminate disc mostly absent, if
present comprising a single central structure ..... 4
3. Staminate flowers having 3(-4) sepals, 5-10
disc segments, and 6-20 stamens; leaf blades
6-16 cm long, unlobed; pollen inaperturate .
....... ........................ . section B ia
3. Staminate flowers having 5 sepals, no disc, and
ca. 40 stamens; leaf blades 12-25 cm long,
unlobed to shallowly 3-lobed; pollen tricolpate
............................ section Zuckertia
4. Pollen tricolpate, exine baculate, aperture mem-
brane bearing few scattered islands of sexine or
sometimes islands absent; stamens (1 -)3-5(-22),
staminate sepals 3(-5) .......... section Tragia
4. Pollen weakly triporate, exine tectate-punctate,
aperture membrane densely covered with nu-
merous small islands of sexine; stamens 2, sta-
minate sepals 4-5(-6) ...... section Leptobotrys
Tragia section Monadelphae L. J. Gillespie, sect.
nov. TYPE: Tragia guayanensis L. J. Gilles-
pie.
Suffrutices scandentes. Inflorescentia unisexualis ra-
cemosa. Sepala masculina 5; discus nullus; stamina mon-
adelpha, columna staminalis elongata, antheris ad apicemn
aggregatis; pollen 5-colpatum in tectis foveolatum. Sepala
feminea 6 integra; styli elongati basi connati.
Scandent subshrubs. Inflorescences unisexual,
racemose. Staminate sepals 5; disc absent; stamens
monadelphous, staminal column elongate with an-
thers clustered together at apex; pollen 5-colpate,
tectum foveolate. Pistillate sepals 6, entire; styles
elongate, connate at base. Monotypic, T. guaya-
nensis.
Tragia guayanensis L. J. Gillespie, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Rio
Casiquiare entre la boca del Siapo y el caiio
Momoni, 18 Feb.-4 Mar. 1986, Stergios &
Aymard 9182 (holotype, MO staminate; iso-
type, NY pistillate). Figures 1, 2, 5, 7, 9.
Suffrutex scandens. Caules juvenes et petioli longihir-
suti trichomatibus urentibus brevissioribus sparsim obsiti.
Folia elliptica vel obovata sparsim hirsuta basi anguste
cordata. Inflorescentiae unisexuales racemosae; inflores-
centia masculina axillaris; inflorescentia feminea termin-
alis sed a foliis opposita videtur. Sepala masculina 5; discus
nullus; stamina monadelpha; columna staminalis elongata
antheris circa 5 ad apicem aggregatis; pollen 5-colpatum
in tectis foveolatum. Sepala feminea 6 integra; styli elon-
gati cylindrici recti basi connati. Capsula trilobata tri-
chomatibus urentibus obsita.
Climbing subshrub, monoecious?; young stems
slender, hirsute with trichomes 1-2 mm long and
scattered, much shorter urticating trichomes. Leaves
alternate, simple; stipules narrowly triangular or lan-
ceolate, ca. 5 x 2 mm, caducous; petioles 2-5.5
cm long, hirsute and often puberulous; blades char-
taceous, elliptic or ovate-elliptic, 12-17 x 6-12
cm, apex acuminate with acumen ca. 1 cm long,
base narrowly cordate with sinus 0.8-1.6 cm deep,
margin irregularly serrulate with tooth apex mi-
nutely glandular, sparsely hirsute on both surfaces,
midrib puberulous and often more densely hirsute
on upper surface; venation pinnate, 3-nerved or
obscurely 5-nerved at base, secondary veins mostly
subopposite in 5-7 pairs with basal pair opposite at
blade base, tertiary veins weakly percurrent, qua-
ternary veins reticulate; petiolar and laminar glands
absent. Inflorescence slender, racemose, unisexual,
flowers borne singly in bract axil; axes puberulous;
bracts lanceolate or narrowly lanceolate; staminate
inflorescence ca. 3-5.5 cm long, axillary, bracts 1-
2.5 mm long; pistillate inflorescence (known only in
fruiting stage) ca. 24 cm long, terminal but ap-
pearing leaf-opposed, bracts ca. 3 mm long. Sta-
minate pedicel 3-3.5 mm long, sparsely puberulous;
bud narrowly oblong, 2-4 mm long, broadly acute
or obtuse at apex; sepals 5, very narrowly oblong,
3-4 mm long, ca. 1 mm wide, acuminate at apex,
valvate, sparsely puberulous, reflexed with margins
revolute at anthesis; corolla and disc absent; stamens
apparently 5, monadelphous; filaments connate into
an elongate staminal column, 2.5-3.1 mm long, ca.
0.4 mm diam., bearing a dense cluster of ca. 5
anthers, 1.1-1.5 mm diam., ca. 0.7-0.9 mm high;
pollen oblate-spheroidal to suboblate, 45-52 hem in
equatorial diameter, 40-49 �m in polar diameter,
5-colpate, colpus with uneven margins, exine tec-
tate-perforate, ca. 2 �m thick, tectum foveolate and
microverrucate. Pistillate pedicel ca. 1 mm long,
Gillespie
Tragia
331

Novon
5
Figures 1-6. Pollen and trichome morphology of Tragia. 1, 2. SEM of pollen of T guayanensis (Stergios &
Aymard 9182, MO). -1. Polar view showing the five colpi. -2. Close-up of mesocolpium and two colpi. 3, 4.
SEM of pollen of T. tabulaemontana (Granville 3570, U). -3. Polar view. -4. Close-up of colpus with sexine
fragment or island on apertural membrane. -5. SEM of leaf blade adaxial surface of T. guayanensis (Williams
14990, US) showing hirsute and puberulous midrib. -6. SEM of inflorescence axis of T. tabulaemontana (Granville
332
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Volume 4, Number 4
1994
densely puberulous (pistillate flower description based
on old flowers on infructescence axis); sepals 6,
ovate, 2.8-3.8 mm long, attenuate at apex, dis-
tinctly imbricate, entire-margined, sparsely pubes-
cent at apex and along margin; corolla and disc
absent; ovary 3-lobed, ca. 0.8-0.9 x 1.3-1.4 mm,
3-locular with 1 ovule per locule, densely covered
with urticating trichomes; styles 3, mostly free, 6-
10 mm long, 0.5-0.8 mm diam., cylindrical, straight,
connate for 1-2.5 mm of length, papillose at apex.
Fruiting pedicel 3-6 mm long; sepals persistent,
mostly reflexed; capsule 3-lobed (irregularly so if
fewer than 3 seeds), ca. 5-5.5 mm long, dehiscing
into 3 bivalved mericarps, each mericarp ca. 5-5.5
mm long and thick, 6.4-6.8 mm wide; pericarp
woody, ca. 0.4-0.7 mm thick, sparsely covered with
urticating trichomes; columella 2.5-3 mm long, per-
sistent, with 3 perpendicular apical arms 1-1.5 mm
long; seeds 3, subglobose, 4.5-4.9 mm diam., pale
brown with paler branched striations and darker
brown markings, inner surface somewhat obtusely
angular.
Distribution. Known only from lowland rainforest
of the upper Orinoco Basin and Rio Casiquiare in
Amazonas, Venezuela.
While considered as belonging to Tragia on the
basis of the combination of slender, entire, mostly
free styles, glabrous staminate receptacle, twining
habit, and urticating trichomes, T. guayanensis is
unique in that genus in having 5-colpate pollen (Figs.
1, 2) and an elongate staminal column (Fig. 7b).
Pollen of all other examined species of Tragia is
3-colpate, weakly 3-aperturate, or inaperturate (Gil-
lespie, 1994), with the exception of T. rubiginosa
Huft, which is 4-colpate (pers. obs.). Tragia typically
has stamens that are free or fused only at their base
(e.g., Fig. 8d). Several species have connate or partly
connate stamens, but none have an elongate staminal
column; for example, T nigricans Bush (sect. Tra-
gia) has filaments connate for one-third to one-half
of their length (Miller & Webster, 1967) and T.
scandens (Baillon) Mueller Argoviensis (monotypic
section Lassia of Madagascar) has stamens connate
into a very short, broad disclike structure (Baillon,
1858: pl. 4, figs. 24, 25; pers. obs.).
Tragia guayanensis is a distinct species of Tra-
gia, apparently not closely related to any other.
While the vast majority of New World Tragia spe-
cies can be easily placed to section, T. guayanensis,
along with T. rubiginosa of Amazonian Peru and
T. biflora Urban of Hispaniola, cannot. These three
species share unisexual, racemose inflorescences (with
staminate inflorescences axillary and pistillate ones
terminal but appearing leaf-opposed), while all other
New World species have bisexual inflorescences
(some African species also have unisexual racemose
inflorescences, but these appear distantly related
based on other floral characters such as pinnatifid
pistillate sepals). Tragia guayanensis and T. rubi-
ginosa also share multicolpate pollen, but differ con-
siderably in floral morphology indicating that the
two species are probably not closely related. Tragia
rubiginosa is distinguished by its sessile anthers and
broad subsessile stigmas and is unique in Tragia in
lacking distinct styles. Huft (1989) suggested a pos-
sible relationship with T. biflora, a species considered
by Liogier (1971) to be intermediate between Tragia
and Platygyna. One or two new sections will likely
be necessary to accommodate T. rubiginosa and T.
biflora if they are retained within Tragia.
Species of Tragia are few and rare in the Amazon
and upper Orinoco basins, but the recent discoveries
of T. guayanensis and T. rubiginosa indicate that
this area harbors some of the most unusual and
phylogenetically interesting species of the genus.
Paratypes. VENEZUELA. Amazonas: rapids of
Trapichote, Delta of Ventuari, 124 m, 21 Apr. 1942,
Williams 14990 (F, US).
Tragia (sect. Tragia) tabulaemontana L. J. Gil-
lespie, sp. nov. TYPE: French Guiana: Sommet
Tabulaire, zone centrale, versant ouest, 600
m, ca. 40 km SE de Saiil, 28 Aug. 1980,
Granville 3637 (holotype, US; isotype, CAY).
Figures 3, 4, 6, 8, 10.
Herba volubilis. Caules petioli et axes inflorescentiae
trichomatibus glanduliferis longistipitatis et trichomatibus
urentibus brevioribus obsiti. Folia oblanceolata vel anguste
elliptica sparsim hirsuta basi anguste cordata. Inflores-
centia racemosa bisexualis basi flore femineo unico. Sepala
masculina 3; stamina 3 apicem versus tenuia basi valde
dilatata. Sepala feminea 6 integra; ovarium trichomatibus
urentibus et glanduliferis longistipitatis dense obsitum; styli
laeves recurvi longitudine 2/% connati.
Twining vine, monoecious; indumentum of long-
stipitate glandular trichomes 0.5-1 mm long, urti-
cating trichomes 0.2-0.5 mm long, and simple tri-
chomes; stems slender, puberulous, sparsely hirsute
with scattered glandular and urticating trichomes.
Leaves alternate, simple; stipules triangular, 2-3
mm long; petiole 0.8-3.2 cm long, puberulous, hir-
4--
3576. CAY) showing puberulous axis with long-stipitate glandular hairs and shorter urticating hairs; note base of
bract at right. Scale bar = 10 im in Figures 1-4, bar = 0.2 mm in Figures 5, 6.
Gillespie
Tragia
333

334
Novon
lmm


Volume 4, Number 4
1994
sute with scattered glandular and urticating tri-
chomes; blade thin-chartaceous, 5-14 x 2-5.5 cm,
narrowly obovate or narrowly elliptic, apex acumi-
nate with acumen 5-12 mm long, base narrowly
cordate with sinus 2-6 mm deep, margin serrate
with serration apex obtuse and minutely glandular,
sparsely hirsute on both surfaces with trichomes ca.
1 mm long, major veins puberulous on upper sur-
face; venation pinnate, secondary veins in 5-7 pairs,
irregularly semicraspidodromous, alternate or sub-
opposite on each side of midrib with basal pair op-
posite diverging at a more acute angle than upper
veins, tertiary and quaternary veins reticulate; pet-
iolar and laminar glands absent. Inflorescence slen-
der, racemose, 4-9 cm long, bisexual, terminal and
appearing leaf-opposed or terminal on short shoots;
axes puberulous, with numerous glandular trichomes
and few scattered urticating trichomes; peduncle
1.3-2.2 cm long, single pistillate flower at basal
node, staminate flowers numerous above, 1 per node;
bracts narrowly triangular-ovate, 1-1.4 mm long,
with scattered glandular trichomes, hirsute at apex.
Staminate pedicel 1-1.7 mm long, puberulous; bud
broadly ovoid, ca. 1 mm long, obtuse at apex; sepals
3, very broadly ovate, 1-1.2 mm long and wide,
sparsely hirsute particularly along margin and at
apex, sometimes with glandular trichomes; corolla
absent; disclike structure obtuse-triangular or 3-lobed
in outline, intrastaminal, adnate to dilated base of
stamens; stamens 3; filaments 0.5-0.6 mm long,
slender and curved upwards at apex, highly dilated
at base; anthers 0.2-0.3 mm long, latrorse; pollen
oblate-spheroidal, 33-38 um in equatorial diameter,
28-33 Am in polar diameter, tricolpate, colpus with
uneven margins and islands of sexine on apertural
membrane, exine intectate-baculate with baculae of-
ten coalesced forming irregularly shaped islands,
surface microverrucate. Pistillate pedicel ca. 1 mm
long, puberulous and with glandular trichomes; se-
pals 6, lanceolate, ca. 2 mm long, puberulous, hir-
sute at apex, with glandular trichomes along margin;
ovary 3-lobed, ca. 1 x 1.8 mm, 3-locular with 1
ovule per locule, densely covered with urticating
and glandular trichomes; styles 3, connate ca. two-
thirds of length into column, 1.7-2 mm long; style
arms 1-1.4 mm long, recurved, with smooth stig-
matic surface. Fruiting pedicel 2-6 mm long; sepals
persistent, reflexed; capsule 3-lobed, ca. 4.5 x 7-
8 mm, dehiscing into 3 bivalved mericarps, each
mericarp 4-4.5 mm long and thick, 6-7.5 mm wide;
pericarp woody, ca. 0.3-0.6 mm thick, sparsely
covered with urticating trichomes and sometimes
glandular trichomes; columella ca. 3 mm long, per-
sistent, with 3 perpendicular apical arms 2-2.4 mm
long. Seeds 3, subglobose, 3.8-4 mm diam., pale
dull yellow with brown or yellowish brown blotches.
Distribution. Known only from Sommet Tabu-
laire in French Guiana, where it has been collected
in submontane forest on the southern and western
slopes.
Tragia tabulaemontana belongs to the neotrop-
ical section Tragia, which is distinguished by rac-
emose bisexual inflorescences, entire pistillate sepals,
tricolpate pollen with an unusual baculate sexine
(Figs. 3, 4; Gillespie, 1994: figs. 64-66), and usually
three staminate sepals and stamens. Relationships
among taxa within this large section are poorly known.
Tragia tabulaemontana appears to belong to a group
that includes T. chlorocaulon Baillon, T. friesii Pax
& K. Hoffmann, T. karsteniana Pax & K. Hoff-
mann, T. mexicana Mueller Argoviensis, and T.
tristis Mueller Argoviensis and which corresponds
approximately to Mueller's section Ratiga (which
is included within sect. Tragia by Miller & Webster
(1967), Mulgura & Gutierrez (1989), and Gillespie
(1994)). This species group is characterized by hav-
ing conical stamens that are slender above and highly
dilated at their base, a smooth stylar stigmatic sur-
face, glandular trichomes, and unlobed leaves. Tra-
gia tabulaemontana may be distinguished by having
oblanceolate or narrowly elliptic leaves that are nar-
rowly cordate at their base (Fig. 10) and long-stip-
itate glandular trichomes on petioles, inflorescence
axes, pistillate sepals, and ovaries (Fig. 6).
The small triangular structure found between the
stamens (Fig. 8d) appears to be an intrastaminal
disc, although its exact nature is uncertain. Similar
structures are found in other species of section Tra-
gia and have been interpreted as either pistillodes
(Pax & Hoffmann, 1919; Miller & Webster, 1967)
or nectaries (Gutierrez & Mulgura, 1986). Inter-
pretation as a disc is preferred since the structures
appear glandular and possibly nectariferous in some
species (e.g., T. tristis) and, while discs or disc
segments are found elsewhere in tribe Plukenetieae,
Figure 7. Tragia guayanensis L. J. Gillespie. -A. Habit showing staminate inflorescence. -B. Staminate flower.
-C. Habit showing infructescence. -D. Pistillate flower. -E. Mericarp of dehisced capsule with enclosed seed. -
F. Seed, lateral view. (A, B based on Stergios & Aymard 9182, MO; C-F based on Stergios & Aymard 9182,
NY.)
Gillespie
Tragia
335


336 Novon
1 Im m 1m
ARnaerini 9q G
Figure 8. Tragia tabulaemontana L. J. Gillespie. -A. Habit. -B. Leaf blade adaxial surface. -C. Upper part
of inflorescence with single open staminate flower with staminate buds above. -D. Staminate flower. -E. Pistillate
flower. -F. Columella and sepals persistent on infructescence axis following capsule dehiscence. -G. Mericarp of
dehisced capsule with enclosed seed. -H. Seed, ventral view. (A-E based on Granville 3637, US; F based on
Granville 3576, CAY; G and H based on Granville 3637, CAY.)


Volume 4, Number 4
1994
Gillespie
Tragia
10
1 cm
Figures 9, 10. Leaf architecture of Tragia (leaf X-rays). -9. T. guayanensis (Williams 14990, US). -10. T
tabulaemontana (Granville 3576, US).
337
obvious pistillodes are not. Tragia tabulaemon-
tana and the widespread weedy species T. volubilis
L. are the only species of section Tragia known
from the Guayana Region including French Guiana.
The section is most diverse in lowland dry tropical
or subtropical regions of the New World and few
species are known from wet tropical regions. Tragia
tabulaemontana is unusual in being apparently re-
stricted to moist or wet submontane forest.
Paratypes. FRENCH GUIANA. Sommet Tabulaire,
versant sud, ca. 50 km SE de Saiil, 550 m, 23 Aug.
1980, Granville 3576 (CAY, U, US).


Novon
KEY TO THE SPECIES OF TRAGIA IN THE GUAYANA REGION
1. Inflorescence unisexual, racemose; staminate
sepals 5; stamens ca. 5, filaments connate into
a central elongate column bearing a cluster of
sessile anthers (sect. Monadelphae) ........
.... ................... . ..T. guayanensis
1. Inflorescence bisexual, racemose or branched;
staminate sepals 3(-4); stamens 1 3 or 8-17,
free or connate only at the base ............. 2
2. Stamens 7-17; inflorescence appearing dichot-
omous, consisting of staminate main axis to 18
cm long and a pistillate basal branch to 10 cm
long; capsule pedicel 1-4 mm long (sect. Bia)
2. Stamens 1-3; inflorescence racemose, to 10 cm
long, with 1 (or 2) pistillate flower(s) at the basal
node(s); capsule pedicel 1-40 mm long (sect.
Tragia) .................................. 4
3. Leaf blade palmately veined, base cordate; sta-
mens 7-10 ...... T. fendleri Mueller Argoviensis
3. Leaf blade pinnately veined, base acute or ob-
tuse; stamens 14-17 .................. .
...... T. lessertiana (Baillon) Mueller Argoviensis
4. Glandular trichomes present on stems, petioles,
and inflorescences; inflorescence 4-9 cm long;
capsule pedicel less than 0.5 cm long ......
............ . T. tabulaemontana
4. Glandular trichomes absent; inflorescence 1-4
cm long; capsule pedicel 1.5-4 cm long ....
.................. . ...... . . T volubilis
Acknowledgments. I thank Kenneth Wurdack
for locating specimens of T. guayanensis and rec-
ognizing them as an undescribed and unusual species
of Tragia. Research was supported by the Smith-
sonian Institution's Biological Diversity of the Guian-
as Program. This study is publication no. 13 in the
series "Biological Diversity of the Guianas."
Literature Cited
Baillon, H. E. 1858. Etude generale du groupe des
Euphorbiacees. Victor Masson, Paris.
Gillespie, L. J. 1993. Euphorbiaceae of the Guianas:
Annotated species checklist and key to the genera.
Brittonia 45: 56 94.
. 1994. Pollen morphology and phylogeny of
the tribe Plukenetieae (Euphorbiaceae). Ann. Mis-
souri Bot. Gard. 81: 317-348.
Gutierrez de Sanguinetti, M. M. & M. E. Mulgura de
Romero. 1986. Una nueva especie de Tragia (Eu-
phorbiaceae). Darwiniana 27: 491-497.
Huft, M. J. 1989. New and critical taxa of Euphorbi-
aceae from South America. Ann. Missouri Bot. Gard.
76: 1077-1086.
Liogier, A. H. 1971. Novitates Antillanae IV. Euphor-
biaceae. Mem. New York Bot. Gard. 21: 123-133.
Miller, K. I. & G. L. Webster. 1967. A preliminary
revision of Tragia (Euphorbiaceae) in the United
States. Rhodora 69: 241-305.
Mori, S. A. 1991. The Guayana Lowland Floristic Prov-
ince. Compt. Rend. Sommaire Seances Soc. Bio-
geogr. 67: 67-75.
Mulgura de Romero, M. E. & M. M. Gutierrez de San-
guinetti. 1989. Actualizaci6n taxon6mica de Tra-
gia (Euphorbiaceae) para Argentina y regiones lim-
itrofes. Darwiniana 29: 77-138.
Pax, F. A. & K. Hoffmann. 1919. Euphorbiaceae-
Plukenetiinae. In: A. Engler (editor), Das Pflanzen-
reich IV.147.XI.(Heft 68): 1 108. W. Engelmann,
Leipzig.
338