SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 86 

A Contribution to the Guianan 

Flora: Dalechampia, Haemutostemon, 

Omp ha lea, Peru, Plukene tia, 

and Tragia (Euphorbiaceae) 

with Notes on Subfamily Acalyphoideae 

Lynn J. Gillespie 

and W Scott Armbruster 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1997 
ABSTRACT 

Gillespie, Lynn J., and W. Scott Armbruster. A Contribution to the Guianan Flora: 

Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, and Tragia (Euphorbiaceae) with 

Notes on Subfamily Acalyphoideae. Smithsonian Contributions to Botany, number 86, 48 

pages, 14 figures, 1997.-Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, and 

Tragia (Euphorbiaceae, subfamily Acalyphoideae) are treated for the Guianas. Notes on 

Guianan Euphorbiaceae, subfamily Acalyphoideae, tribes Omphaleae, Pereae, and Plukenetieae, and ecological notes on each species are given. Also included is a key to the subfamilies 

of Euphorbiaceae and a key to the Guianan genera of subfamily Acalyphoideae. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVER 

DESIGN: Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging-in-Publication Data 

Gillespie, Lynn J. 

A contribution to the Guianan flora : Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, and Tragia 

(Euphorbiaceae) with notes on subfamily Acalyphoideae / Lynn J. Gillespie and W. Scott Armbruster. 

p. cm.-(Smithsonian contributions to botany ; no. 86) 

Includes bibliographical references (p. ) and indexes. 

1. Euphorbiaceae-Guiana-Classification. I. Armbruster, W. Scott. 11. Title. 111. Series. 

QKIS2747 no. 86 

IOK495.E91 

%O s-dc21' 

[583'.69'0988] 97-10535 

CIP 

@ The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials 239.48-1984. 
Contents 

Page 

Introduction ........................................ 1 

Acknowledgments .................................. 1 

EUPHORBIACEAE in the Guianas ............................. 1 

Key to the Subfamilies of Euphorbiaceae in the Guianas ............... 2 

Subfamily ACALYPHOIDEAE .............................. 3 

Key to the Genera of Acalyphoideae in the Guianas ............... 3 

Tribe OMPHALEAE .................................. 6 

1 . Genus Omphalea L ............................... 6 

1 . Omphalea diandra L ............................ 6 

Tribe PEREAE .................................... 8 

2 . Genus Pera Mutis ............................... 8 

Key to the Species of Pera .......................... 9 

1 . Pera bicolor (Klotzsch) Muell . Arg .................... 9 

2 . Pera decipiens (Muell . Arg.) Muell . Arg ................. 11 

3 . Pera glabrata (Schott) Bail1 ........................ 13 

Tribe PLUKENETIEAE ................................ 13 

3 . Genus Dalechampia L ............................. 14 

Key to the Species of Dalechampia ..................... 14 

1 . Dalechampia affinis Muell . Arg ...................... 16 

2 . Dalechampia attenuistylus Armbruster ................. 17 

3 . Dalechampia brevicolumna Armbruster ................. 17 

4 . Dalechampia brownsbergensis Webster & Armbruster ......... 19 

5 . Dalechampia dioscoreifolia Poepp .................... 19 

6 . Dalechampia fragrans Armbruster ................... 19 

7 . Dalechampia heterobractea Armbruster ................. 23 

8 . Dalechampia magnoliifolia Muell . Arg .................. 23 

9 . Dalechampia megacarpa Armbruster .................. 23 

10 . Dalechampia micrantha Poepp ...................... 24 

11 . Dalechampia olympiana Kuhlman & Rodrigues ............ 24 

12 . Dalechampia papillistigma Armbruster ................. 24 

13 . Dalechampia pawibracteata Lanj ..................... 25 

14 . Dalechampia scandens L ......................... 25 

14a . Dalechampia scandens var . scandens ............... 25 

14b . Dalechampia scandens var . fallax Muell . Arg ........... 25 

15 . Dalechampia stipulacea Muell . Arg ................... 27 

16 . Dalechampia tiliifolia Lam ........................ 27 

17 . Dalechampia aff . cissifolia Poepp ..................... 28 

18 . Dalechampia aff . triphylla Lam ...................... 28 

4 . Genus Haematostemon (Muell . Arg.) Pax & K . Hoffm ........... 28 

1 . Haematostemon guianensis Sandwith .................. 30 

5 . Genus Plukenetia L .............................. 30 

Key to the Species of Plukenetia ...................... 31 

31 

2 . Plukenetia polyadenia Muell . Arg .................... 31 

3 . Plukenetia supraglandulosa L.J. Gillespie ................ 33 

1 . Plukenetia loretensis Ule ......................... 

... 

111 
iv SMITHSONIAN CONTRIBUTIONS TO BOTANY 

4 . Plukenetia verrucosa Smith ....................... 35 

5 . Plukenetia volubilis L ........................... 35 

6 . Genus Tragia L ................................ 37 

Key to the Species of Tragia ........................ 37 

1 . Tragia fendleri Muell . Arg ........................ 37 

2 . Tragia lessertiana (Baill.) Muell . Arg .................. 39 

3 . Tragia tabulaemontana L.J. Gillespie .................. 39 

Appendix 1 . Numerical List of Accepted Taxa ..................... 42 

4 . Tragia volubilis L ............................. 41 

Appendix 2 . Collections Studied ............................ 43 

Literature Cited ..................................... 45 

Index to Scientific Names ................................ 47 

Index to Vernacular Names ............................... 48 
A Contribution to the Guianan 

Flora: Dalechampia, Haematostemon 

Omphalea, Pera, Plukenetia, 

and Tragia (Euphorbiaceae) 

with Notes on Subfamily Acalyphoideae 

Lynn J. Gillespie 

and K Scott Armbruster 

Introduction 

This contribution treats three of the 10 tribes of subfamily 

Acalyphoideae known from the Guianas. Tribe Plukenetieae 

includes four genera in the Guianas, whereas the remaining two 

tribes, Omphaleae and Pereae, each comprise a single genus. A 

more concise version of this contribution eventually will be 

published as part of the Euphorbiaceae treatment for the Flora 

ofthe Guianas. However, only a very small part of the Guianan 

Euphorbiaceae currently is under study, and there is no 

scheduled completion date for the family treatment. No further 

work by the two authors on the taxonomy of Euphorbiaceae of 

the Guianas is anticipated at present. Current interest in the 

Guianan flora and the need for up-to-date treatments of 

neotropical taxa make it timely to publish the completed study 

of these tribes at this time. 

This treatment is limited to the three Guianan countries, 

Guyana, Surinam, and French Guiana. Species from neighboring areas within the Guayana Region also are included or 

mentioned if they are thought likely to occur in the Guianas. 

The Guayana Region (Gillespie, 1993a; equivalent to the 

Guayana Lowland Floristic Province as defined by Mori, 1991) 

is a more biogeographically natural region than the Guianas 

and includes, in addition to the Guianas, the Venezuelan 

Guayana (comprising the states of Amazonas, Bolivar, and the 

Delta Amacuro) and Amazonian Brazil north of the Amazon 

and Negro rivers. 

In addition to the floristic taxonomic treatments of the genera 

Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, 

and Tragia, we have included notes on Guianan Euphorbiaceae, subfamily Acalyphoideae, and tribes Omphaleae, 

Pereae, and Plukenetieae. Also included are keys to the five 

subfamilies of Euphorbiaceae and to the 18 genera of 

Acalyphoideae known from the Guianas. 

ACKNOWLEDGMENTS.-we acknowledge the support of the 

Biological Diversity of the Guianas Program, Department of 

Botany, National Museum of Natural History, Smithsonian 

Institution. Cathy Pasquale and Alice Tangerini are thanked for 

providing illustrations. We acknowledge the Flora of the 

Venezuelan Guayana Project, Missouri Botanical Garden, for 

providing Figure ~A,B,E by Bruno Manara. W.S. Armbruster 

acknowledges support by US National Science Foundation 

grants BSR-850903 1 and BSR-9006607 and thanks Ruth 

Stafford for curatorial assistance. This contribution is publication 20 in the series “Biological Diversity of the Guianas,” 

published by the Biological Diversity of the Guianas Program 

(address above). 

Lynn J. Gillespie, Department of Botany, National Museum of 

National Histoy, Smithsonian Institution, Washington D. C. 20560, 

and the Research Division, Canadian Museum of Nature, Box 3443, 

Station D, Ottawa, Ontario KIP 6P4, Canada; W Scott Armbruster, 

Department of Biology and Wildrife and Institute of Arctic Biologv, 

University of Alaska Fairbanks, Fairbanks, Alaska, 99775-7000. 

EUPHORBIACEAE in the Guianas 

The Euphorbiaceae is considered to be the fifth largest 

flowering plant family and comprises about 300 genera and 

7500 species. Forty-eight genera and 17 1 species are known to 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

be indigenous or extensively naturalized in the Guianas, 

whereas an additional six genera and 22 species are known only 

in cultivation or are naturalized to a very limited extent in the 

Guianas (Gillespie, 1993a; additions to checklist listed below). 

Twenty-one species (12%) of Guianan Euphorbiaceae are 

strictly endemic to the Guianas, whereas 55 species (32%) are 

endemic to the more broadly defined Guayana Region 

(Gillespie, 1993a, with additional species listed below). 

Five new species have been described from the Guianas 

since publication of the most recent checklist of Guianan 

Euphorbiaceae (Gillespie, 1993a). These are Dalechampia 

brevicolumna Armbruster, D. heterobractea Armbruster 

(Armbruster, 1996), Mabea salicoides Esser (Esser, 1993), 

Plukenetia supraglandulosa L.J. Gillespie (Gillespie, 1993b), 

and Tragia tabulaemontana L.J. Gillespie (Gillespie, 1994b). 

New records for the Guianas include Croton draconoides 

Muell. Arg. (Hoff et al., 1996), Dalechampia olympiana 

Kuhlman & Rodrigues (Armbruster, 1996), Manihot aff. 

quinquepartita Rogers and Appan (Gillespie, unpublished 

data), Phyllanthus caribeus Urb. (Webster, pers. comm., 1993), 

and Senefelderopsis croizatii Steyerm. (Esser, 1995). Additionally, three species of Dalechampia, D. stipulacea Muell. Arg., 

D. aff. cissifolia, D. aff. triphylla, that are new to the Guianas 

are presented here. 

Previous floras and checklists of Euphorbiaceae of the 

Guianas include The Euphorbiaceae of Surinam (Lanjouw, 

193 1; additions by Gorts-van Rijn, 1976), Flore de la Guiane 

Francaise (Lemee, 1952), and Checklist of Woody Plants of 

Guyana (Mannega et al., 1988). The series Botany of the 

Guayana Highland (Jablonski, 1965, 1967) treats the Euphorbiaceae flora of the region adjacent to the western border of 

Guyana, as will the Flora of the Venezuelan Guayana 

(Steyermark et al., 1995; Webster et al., in prep). Additional 

references, including recent revisionary studies of neotropical 

Euphorbiaceae, are listed in Gillespie (1 993a). 

Five subfamilies of Euphorbiaceae are currently recognized 

(classification of Webster, 1975, 1994). The family previously 

was divided into two major groups based on the number of 

ovules per locule, the biovulate taxa belonging to the 

Phyllanthoideae sensu Pax and the uniovulate taxa to the 

Crotonoideae sensu Pax (Pax, 1890; Pax and Hoffmann, 193 1; 

for an historical account refer to Webster, 1987). In Webster’s 

classification two subfamilies, Phyllanthoideae and Oldfieldioideae, include all biovulate taxa, whereas the remaining 

three subfamilies, Acalyphoideae, Crotonoideae sensu Webster, and Euphorbioideae, comprise all uniovulate taxa. 

All five subfamilies of Euphorbiaceae occur in the Guianas. 

The largest subfamily in the Guianas (excluding cultivated 

genera and species) is the Acalyphoideae (17 genera, 54 

species), followed by the Crotonoideae (1 1 genera, 50 species), 

Euphorbioideae (1 1 genera, 35 species), Phyllanthoideae (8 

genera, 3 1 species), and Oldfieldioideae (1 genus, 1 species). 

Key to the Subfamilies of Euphorbiaceae in the Guianas 

1. Ovules paired in each locule of the ovary; fruit with 1-10 (often 6) seeds, 1 or 2 in 

each locule; staminate calyx-lobes imbricate in bud; latex absent; leaf (or leaflet in 

Piranhea) blade pinnately veined, the margins unlobed, entire, or obscurely 

crenate; leaf lacking petiolar glands and usually lacking embedded laminar glands; 

indumentum simple or lepidote ......................... .2 

Ovules solitary in each locule of the ovary; fruit with 1-3 (rarely to 20) seeds, always 

solitary in each locule; staminate calyx-lobes valvate, imbricate, reduced, or 

absent; latex present or absent; leaf blade pinnately or palmately veined, the 

margins often lobed, serrate, or dentate; leaf often with petiolar glands and/or 

embedded laminar glands; indumentum simple, lepidote, stellate, or dendritic 

........................................... 3 

2. Leaves simple; seeds ecarunculate; herbs, shrubs, or trees [Podocalyx of subfamily 

Oldfieldioideae keys out here, but known only from the Guayana Region outside 

of the Guianas] .................. Subfamily PHYLLANTHOIDEAE 

Leaves trifoliate; seeds carunculate; dioecious trees [Piranhea, is the only genus in 

the Guianas.] .................... Subfamily OLDFIELDIOIDEAE 

Staminate bracts biglandular at base (except Hura and Senefeldera) or inflorescence 

cyathial (i.e., flowers contained within a cupular cyathium formed by 5 connate 

bracts); staminate calyx-lobes imbricate, reduced, or absent, usually not covering 

the anthers in bud; petals absent; floral disc (nectary) absent; leaf blades simple, 

pinnately veined, unlobed; indumentum, if present, simple or dendritic (in Mabea 

spp. and Senefeldera spp.); latex milky white, sometimes scant or rarely absent 

3. 

........................... Subfamily EUPHORBIOIDEAE 
NUMBER 86 3 

Staminate bracts eglandular (except Omphalea and Pausandra), inflorescence not 

cyathial; staminate calyx-lobes valvate or imbricate, covering the anthers in bud; 

petals present or absent; floral disc present or absent; leaf blades simple or 

compound, pinnately or palmately veined, lobed or unlobed; indumentum simple, 

stellate, or lepidote; latex clear, colored, milky white, or absent ........ 4 

Petals present at least in staminate flowers (if absent then sepals petaloid or latex 

present); latex milky white, colored, clear, or rarely absent; petiolar glands often 

present; staminate calyx-lobes mostly imbricate in bud; styles bifid or multifid 

(entire in Aleurites and Codiaeum, both cultivated genera), free or connate only at 

base; seeds mostly carunculate; pollen with crotonoid exine sculpture (clavae 

arranged in regular hexagonal pattern) ....... Subfamily CROTONOIDEAE 

Petals absent (present in Caperonia and Pogonophora); latex absent (pinkish to red 

in Omphalea); petiolar glands absent (present in Omphalea); staminate 

calyx-lobes mostly valvate in bud; styles entire to multifid, free to entirely 

connate; seeds mostly ecarunculate; pollen never with crotonoid exine sculpture 

4. 

........ Subfamily ACALYPHOIDEAE 

Subfamily ACALYPHOIDEAE 

The Acalyphoideae, with 116 genera in 20 tribes, is the 

largest and most complex subfamily in the Euphorbiaceae 

(Webster, 1994). Species characteristically lack latex, petiolar 

glands, inflorescence bract glands, and petals, and they 

typically have staminate flowers with valvate sepals. The 

subfamily is not defined by any uniquely derived character or 

character complex, but instead it is characterized primarily on 

the basis of characters considered plesiomorphic in the 

Euphorbiaceae (with the exception of apetalous flowers, a state 

that appears to have evolved several times in the family). As 

currently circumscribed, the Acalyphoideae is paraphyletic, 

comprising all uniowlate taxa that do not fit in either the 

Crotonoideae or Euphorbioideae. Although a large group of 

related genera appears to form the core of the subfamily (e.g., 

tribes Acalypheae and Alchornieae), there are many genera that 

appear to be phylogenetically isolated (e.g., Chaetocarpus 

Thwaites, Clutia L., Omphalea, Peru, and Pogonophora Miers 

ex Benth.). ' 

In Acalyphoideae five tribes are unigeneric, whereas three 

comprise only two genera. The largest tribe is the Acalypheae 

with 30 genera. Eighteen genera (including the cultivated genus 

Ricinus) in 10 tribes of Acalyphoideae are known from the 

Guianas. In addition, the genus Acidoton, known from 

neighboring Bolivar state, Venezuela, may be expected to 

occur in the Guianas and is included in the key to genera. 

DESCRIPTION.-Trees, shrubs, herbs, or vines; monoecious 

or dioecious; latex absent or rarely present and then not milky 

white; indumentum simple, lepidote, or stellate. Leaves simple 

or rarely palmately compound, alternate or rarely opposite, 

usually stipulate; blades pinnately or palmately veined, margin 

entire, serrate, or lobed; petiolar glands absent (present in 

Omphalea), laminar glands often present. Inflorescence various; bracts eglandular (biglandular in Omphalea). Flowers 

lacking corolla (except Caperonia and Pogonophora); disc 

present or absent. Staminate calyx mostly valvate or sometimes 

imbricate, anthers covered in bud; pollen grains 3(-4)- 

colporate or -colpate, rarely porate or inaperturate, exine 

tectate, semitectate, or rarely intectate, never with crotonoid 

exine sculpture. Ovary 3-locular, rarely I-, 2-, or 4-locular, 

ovules solitary in each locule; styles entire to multifid, free to 

entirely connate. Fruit capsular or rarely baccate; seeds 

(1-)3(-4), solitary in each locule, mostly ecarunculate; testa 

mostly dry, sometimes fleshy. 

Key to the Genera of Acalyphoideae in the Guianas 

1. Petals present in both staminate and pistillate flowers; staminate flower with 

pistillode ..................................... 2 

Petals absent from both staminate and pistillate flowers; staminate flower with or 

without pistillode ................................. 3 

2. Leaf blade margins entire, the secondary veins arcuate; stamens 5; dioecious trees 

.................................... Pogonophora 

Leaf blade margins finely serrate, the secondary veins closely set and distinctly 

parallel; stamens 10; monoecious herbs ................ Caperonia 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

3. Inflorescences pseudanthial, the flowers subtended by an involucre of 2 free or 

connate bracts, the involucre usually showy, sometimes reduced ........ 4 

Inflorescences not pseudanthial .......................... 5 

4. Flowers enclosed within globose, petal-like involucre, consisting of 2 connate 

bracts, opening along one side at anthesis; involucres unisexual, in axillary 

clusters on woody branches below leaves; leaves simple, blade not lobed; 

dioecious trees ............................... 2. Peru 

Flowers subtended by bilabiate involucre of 2 palmately veined foliaceous or 

stipuliform bracts; involucres bisexual, axillary or terminal on short shoots; leaves 

simple to palmately compound, blade often lobed; monoecious twining vines or 

rarely subshrubs .......................... 3. Dalechampia 

Stamens 100-1000, the filaments partly and irregularly connate; leaf blade peltate, 

palmately 7- to 1 1 -lobed; inflorescence bisexual, the pistillate flowers numerous 

and distal; seeds carunculate; monoecious, weak-stemmed shrubs, cultivated in 

the Guianas ................................ Ricinus 

Stamens 2-60, filaments free to completely connate; leaf blade not peltate, rarely 

lobed; inflorescence unisexual or bisexual with pistillate flowers proximal; seeds 

mostly ecarunculate ............................... .6 

6. Flowers in axillary glomerules; staminate sepals imbricate in bud; capsule 

spiny-tuberculate; seeds black, shiny, with large red caruncule; dioecious shrubs 

or small trees ............................ Chaetocarpus 

Flowers in spicate, racemose, or paniculate inflorescences (terminal glomerules in 

Bernardia); staminate sepals valvate in bud (imbricate in Omphalea); capsule not 

spiny-tuberculate; seeds ecarunculate or rarely with very small caruncle, usually 

not black. .................................... 7 

7. Androecium mushroom-shaped, the stamens 2, completely connate, with connective greatly expanded; staminate sepals imbricate in bud; inflorescence terminal, 

paniculate, the cymules bisexual or staminate; bracts usually large, foliaceous, 

Androecium not mushroom-shaped, the stamens 2-60, free or connate only at base, 

connective not or only slightly expanded; staminate sepals valvate in bud; 

inflorescence axillary or terminal, spicate or racemose, the cymules unisexual or 

flowers solitary at a node; bracts small, not foliaceous (large sessile bracts often 

subtend pistillate flower(s) in Acalypha); latex absent or rarely present and then 

not colored. ................................... 8 

Styles entire, basally to completely connate, often massive; inflorescence usually 

bisexual with 1 to several basal pistillate flowers; plants often twining vines (if not 

then urticating hairs present or styles completely connate and massive), sometimes 

5. 

petiolate; latex red, sometimes scant; monoecious lianas ...... 1. Omphalea 

8. 

with urticating hairs, monoecious (dioecious in Acidoton) ............ ............................... 9 [Tribe Plukenetieae] 

Styles branched or entire, free or sometimes basally connate (rarely long-connate 

and then bifid), not massive, rarely absent; inflorescence unisexual (sometimes 

bisexual in Acalypha and Bernardia and then the styles branched); plants not 

twining vines, lacking urticating hairs, dioecious (usually monoecious in 

9. Ovary 4-locular; fruit 4-lobed or subglobose, dehiscent or indehiscent; leaf blade 

with circular laminar glands at base; twining vines or lianas lacking urticating 

hairs ................................. 5. Plukenetia 

Ovary 3-locular; fruit 3-lobed, dehiscent; leaf blade lacking laminar glands; habit 

10. Urticating hairs absent; sepals 4; stamens 4; styles completely connate forming a 

massive hollow cup-shaped structure; leaf blades pinnately veined; monoecious 

shrubs or trees ......................... 4. Haematostemon 

Acalypha and Bernardia) ............................ 12 

various, urticating hairs present or absent .................... 10 
NUMBER 86 5 

Urticating hairs present at least on ovary and capsule; sepals 3 or 5; stamens (I-)3 

or numerous; styles slender, only partly connate; leaf blades pinnately or 

1 1. Anthers with apical tuft of urticating hairs; leaves glabrescent, pinnately veined; 

dioecious shrubs or trees ......................... Acidoton 

Anthers not apically tufted; leaves with urticating hairs or rarely glabrescent, 

pinnately or palmately veined; monoecious twining vines (in the Guayana Region) 

Styles laciniate (i-e., multifid with many filiform segments); pistillate bracts usually 

large, conspicuous, and lobed; anthers vermiform (elongate and worm shaped); 

palmately veined ................................. 11 

...................................... 6. Tragia 

inflorescence spicate; herbs, shrubs, or small trees, mostly monoecious ..... 

...................................... Acalypha 

12. 

Styles entire or bifid, not laciniate; pistillate bracts small and inconspicuous; anthers 

not vermiform, usually globose or ellipsoid; inflorescence various; habit various, 

mostly dioecious ................................. 13 

Pistillate or bisexual inflorescence glomerate, terminal; staminate inflorescence, if 

present, spicate or racemose, shorter than 5(-6) cm; stamens 4-6, the anthers 

constricted at middle, having the appearance of 4 globose locules; styles stout, 

short, free, bipartite to base; leaf blade shorter than lo(-12) cm, usually hirsute, 

Pistillate and staminate inflorescences spicate, racemose, or paniculate, longer than 

5 cm, axillary or terminal; stamens 2-60, the anthers not constricted; styles 

various; leaf blade usually longer than 10 cm, rarely hirsute, the margins entire to 

sermlate; shrubs or trees ............................. 14 

Stamens 15 or more; styles bifid; capsules ovoid or pear-shaped, mostly circular in 

cross section, greater than 1.5 cm in diameter; leaf blades palmately or pinnately 

veined; hairs minutely stellate .......................... 15 

Stamens fewer than 10; styles entire, indistinctly bifid, or absent; capsules 

subglobose to oblate, lobed in cross section, less than 1.5 cm in diameter; leaf 

blades pinnately veined (either 1 or 3-veined at base); hairs simple or minutely 

stellate ...................................... 16 

15. Carpels 2, styles 2; pistillate calyx cup-shaped, only slightly lobed; staminate 

inflorescence axillary; all stamens fertile; pistillode present; leaf blade narrowly 

obovate. ................................ Gavarretia 

Carpels 3, styles 3; pistillate calyx deeply lobed into sepals; staminate inflorescence 

terminal; inner stamens often sterile; pistillode absent; leaf blade cordiform, ovate, 

or elliptic ............................... Conceveiba 

Styles absent (i.e., stigmas subsessile); stamens 2(-3); leaf blades 1-veined and 

narrowly cuneate at base, eglandular ............... Adenophaedra 

Styles present; stamens 3-8; leaf blades 1- or 3-veined at base, acute to rounded at 

base, usually with laminar glands ........................ 17 

17. Carpels 2; styles 2; elongate, slender, free; stamens usually 8, pistillode absent; 

hairs minutely stellate ......................... Alchornea 

Carpels 3; styles 3, short, free or connate at base; stamens 3-8, pistillode present or 

absent (if stamens 8, then pistillode present); hairs simple ........... 18 

18. Staminate flower with large pubescent pistillode, 5-8 stamens, the anthers 

apiculate; pistillate flowers subsessile; capsule -5 mm in diameter; outer seed coat 

fleshy, bright red; leaf blades less than 15 cm long, distinctly 3-veined and without 

stipels at base ............................ Alchorneopsis 

Staminate flower without pistillode, with 3-5 stamens, the anthers not apiculate; 

pistillate flowers with pedicel longer than 4 mm; capsule -1 cm in diameter; outer 

seed coat dry, brownish; leaf blades usually longer than 15 cm, distinctly I-veined 

and with pair of stipels at base ................... Aparisthmium 

13. 

the margins coarsely serrate; herbs or weak-stemmed shrubs ..... Bernardia 

14. 

16. 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Tribe OMPHALEAE 

Tribe Omphaleae (Pax & Hoffm.) Webster, is anomalous in 

the Acalyphoideae in having latex, biglandular petioles, usually 

biglandular inflorescence bracts, and imbricate sepals. The 

tribe shares the above characters with subfamily Euphorbioideae, but it differs in having palmately veined, sometimes 

lobed leaves, and in having pinkish to red rather than milky 

white latex. With the exception of biglandular bracts, the tribe 

shares the same characters with the Crotonoideae, but it does 

not have pollen with a crotonoid exine sculpture, a diagnostic 

character of that subfamily (known to be absent in only a single 

species of Micrandru; Nowicke, 1994). This heterogenous mix 

of subfamilial characters and the lack of any closely related 

taxa indicates that the Omphaleae most likely represent a rather 

ancient lineage within the uniovulate euphorbs. A single genus 

comprises tribe Omphaleae. 

Omphalea is distributed pantropically, except for most of 

mainland Africa, with centers of diversity and endemism in 

Madagascar and the Caribbean. This peculiar, somewhat 

relictual distribution may perhaps be further evidence of its 

antiquity. Omphalea may be distinguished by its unusual 

mushroom-shaped androecium, pinkish to red latex, foliaceous 

inflorescence bracts, and usually large paniculate inflorescences of bisexual or staminate cymules. 

1. Genus Omphalea L. 

Omphalea L., Syst. nat., ed. 10, 2:1264, 1759. [Qpe: Omphalea triandra L.] 

DESCRIPTION.-Trees, shrubs, or lianas, monoecious; latex 

red or pink, scant to copious; indumentum of simple hairs. 

Leaves simple, petiolate; stipules small, deciduous; blade 

margin entire to deeply lobed, pinnately or palmately veined; 

pair of glands present at petiole apex or blade base; laminar 

glands usually scattered on abaxial surface of blade. Inflorescence a terminal thyrse, appearing spicate, racemose, or 

paniculate; cymose subunits bisexual with central flower(s) 

pistillate or distally unisexual and all staminate; bracts 

subtending cymose subunits large, leaf-like, often biglandular; 

bracteoles small, triangular, usually eglandular. Staminate 

flowers pedicellate; sepals 4-5, imbricate; corolla absent; disc 

extrastaminal, annular, rarely 5-segmented or absent; androecium mushroom-shaped with massive hemispheroidal cap 

formed from highly expanded connate anther connectives, or 

rarely not mushroom-shaped with anthers free and connectives 

not expanded; filaments completely connate into a usually 

slender staminal column or rarely anthers subsessile; anthers 2 

or 3; pistillode absent. Pistillate flowers subsessile to pedicellate; sepals 4-5, imbricate; corolla and disc absent; ovary 

3-locular; styles completely connate, usually massive. Fruit a 

3-seeded schizocarpic capsule or berry; capsule separating into 

3 mericarps, the mericarps fleshy or woody, indehiscent or 

loculicidally dehicent. Seeds globose, ovoid, or ellipsoid, 

ecarunculate, surface smooth, roughened, or tuberculate. 

DISTRIBUTION.-Seventeen species, pantropical; one species 

in the Guianas. 

LITERATURE.-PaX and Hoffmann (1912); Gillespie (1988). 

1. Omphalea diandra L. 

FIGURE 1 

Omphalea diandra L., Syst. nat., ed. 10, 2:1264, 1759. [Type: none 

Omphalea megacaya Hernsley, Hooker’s Icon, pl. 4, pl. 2537, 1897. [TYPE: 

designated.] 

Tobago, Gilloway TRIN 5071 (holotype, K!).] 

DESCRIPTION.-Liana to 30 m or more in forest canopy, 

climbing by means of tendril-like stems; trunk cylindrical, up 

to 20 cm in diameter; growth sympodial. Petiole 4-20 cm long; 

blade chartaceous to subcoriaceous, ovate or elliptic, 15-27 x 

6- 16 cm, apex rounded-acuminate or obtuse-acuminate, base 

shallowly cordate or rounded, glabrescent to velutinous below, 

margin entire, laminar glands scattered near margin below; pair 

of glands present adaxially at petiole apex. Inflorescence a 

paniculate thyrse, 15-65 cm long, cymes condensed, axes 

sparsely to densely pubescent; bracts to 6 cm long, petiolate, 

biglandular, blade oblanceolate or narrow elliptic, to 5 x 0.8 

cm. Staminate pedicel 1-4 mm long; sepals 4 or rarely 5, 

subspherical or broadly ovate, 1.5-3.5 mm long; disc annular; 

androecium mushroom-shaped, staminal column slender, cap 

1-2 mm in diameter, anthers 2. Pistillate pedicel 0.2-2.3 mm 

long; sepals 4 or 5, subspherical or broadly ovate, 1.5-2.5 mm 

long; ovary globose, 2-3.5 mm in diameter, densely pubescent; 

style subglobose, 1.6-2.6 mm in diameter, densely pubescent. 

Fruit subglobose, 8-12 cm in diameter, indehiscent; pericarp 

fleshy leathery, shiny green or greenish yellow. Seeds 3, 

broadly ellipsoid, radially compressed, 3.4-4.8 x 3.3-5 x 

2.2-3.2 cm, surface dark brown, rough, dull; testa 0.5-2 cm 

thick, fleshy, white, not persistent. 

DISTRIBUTION.-Widespread in the Neotropics, from Honduras and the West Indies to Bolivia and Brazil; -160 

collections examined, of which 67 are from the Guianas 

(Guyana: seven collections, Surinam: 34 collections, and 

French Guiana: 26 collections; abbreviated hereafter as G:7, 

S:34, FG:26). 

ECOLOGY.4anopy liana in moist to wet lowland forest, 

below 700 m (below 500 m in the Guianas), often dispersed and 

uncommon, but sometimes locally abundant in riparian forest, 

littoral forest, or at edge of granitic outcrops (e.g., Voltzberg). 

Flowers mostly between June and December in the Guianas. 

Seed germination is epigeal and cryptocotylar, i.e., the 

cotyledons of seedlings are above ground but remain hidden 

within the seed coat (Gillespie, 1988). This is an unusual mode 
NUMBER 86 7 

1 mm -u 1 mm 

FIGURE 1.4rnphalea diandra L.: A, habit, flowering branch; B, cymose partial-inflorescence showing central 

pistillate flower and outer staminate flower buds; C, androecium above annular nectary; D, staminate flower at 

anthesis; E, fruit (A,B,E based on Gent? et al. 30692; C,D based on Gillespie 648). 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

of germination in the Euphorbiaceae, where the majority of 

species have seedlings with cotyledons that emerge from the 

seed coat and are green and photosynthetic. 

SELECTED SPECIMENS ExAMINED.--Guyana: Pomeroon 

River, de la Cruz 3186 (F, GH, MO, NY, UC, US); Groete 

Creek, Lower Essequibo River, Fanshawe 1993 = FD 4729 (A, 

FDG, K, NY, U). Surinam: Beach N of Galibi, mouth of 

Marowijne River, Lanjouw & Lindeman 1045 (NY, U); Lucie 

River, 2- I0 km below confluence of Oost River, Irwin et al. 

55556 (F, K, MO, Ny, U, US, VEN); Voltzberg, Lanjouw 867 

(MO, S, U). French Guiana: Rio Oiapoque, Roche Mon Pere, 

Irwin et al. 48181 (MO, NY, U, UC, US, VEN); Ile de 

Cayenne, Pointe de Bourda, Granville 5753 (U). 

VERNACULAR NAMES.-Surinam: baboenoot, baboenotto, 

idaballie (Arawak), mekoekwaire, sitjo (Carib). French Guiana: 

ouabe, sooke (Taki-Taki), graine de l’anse, liane papaye. 

USES.-Seeds are reported as edible but may be mildly toxic 

unless the embryo is removed. Oil extracted from the seeds has 

been used medicinally as a purgative (Freeman and Williams, 

1927). In the Guianas, necklaces apparently have been made 

out of small polished fragments of the hard seed coat (Duss, 

1897). 

NOTES.-Mature individuals of Omphalea are large canopy 

lianas with one or more cylindrical trunks, whereas young 

individuals are vine-like. The genus is unique among Guianan 

Euphorbiaceae in having a liana habit combined with an 

unusual mode of climbing. Only the distal leafless part of the 

main stem twines and fbnctions in climbing. These distal 

twining stem sections soon abort and frequently appear 

tendril-like following further growth of the axis from a 

sympodial shoot at their base. This method of climbing is in 

contrast to the much more typical method of all other 

Euphorbiaceae vines and lianas in the Guianas (e.g., Dalechampia spp., Plukenetia, and Tragia). These species climb by 

means of twining stems that both bear leaves and continue to 

grow. 

Leaf shape, texture, and pubescence can vary considerably 

depending on maturity of plant and position on plant. Typical 

mature leaves of 0. diandra are ovate, subcordate at base, 

subcoriaceous, and usually pubescent below, whereas juvenile 

leaves are lanceolate, rounded or obtuse at base, membraneouschartaceous, and glabrescent. Leaves from shady understory 

habitats or new lateral shoots from old branches frequently 

resemble juvenile leaves (e.g., Lindeman 5111). 

Tribe PEREAE 

Tribe Pereae (Klotzsch & Gcke.) Pax & K. Hoffm. is unique 

in the Euphorbiaceae in having unisexual pseudanthia consisting of morphologically reduced flowers enclosed by a showy, 

more or less globose, involucral bract. Pseudanthia are 

frequently strongly scented, and the bracts are typically yellow, 

cream, white, or sometimes red in color. Tribe Pereae 

represents one of three independent origins of pseudanthia in 

the Euphorbiaceae; pseudanthia also characterize Dalechampia 

and tribe Euphorbieae (i.e., the cyathium of Euphorbia L. and 

related genera). 

Tribe Pereae includes only the single genus Pera. The highly 

reduced nature of the flowers has to a large degree obscured its 

phylogenetic affinity. Airy Shaw (1973) in recognizing the 

genus as a distinct family, the Peraceae, emphasized its isolated 

position (also discussed by Radcliffe-Smith, 1987). Most 

recently, Webster (1994) suggested that Pera may be most 

closely related to tribe Chaetocarpeae based on similarity in 

pollen, capsule, and seed morphology (e.g., they share 

carunculate seeds, an unusual character in subtribe Acalyphoideae). 

2. Genus Pera Mutis 

Pera Mutis, Kongl. Svenska Vetensk. Acad. Handl., 5:299, t. 8, 1784. [Type: 

DESCRIPTION.-Shrubs or trees; dioecious (or rarely monoecious); indumentum usually of lepidote (flat peltate scale-like) 

or stellate hairs, rarely of simple hairs. Leaves simple, alternate 

or rarely opposite, petiolate, exstipulate, eglandular; blade 

pinnately veined, secondary veins looped, margin entire. 

Inflorescence pseudanthial, unisexual (or rarely bisexual?); 

pseudanthial units pedunculate, fasciculate on short shoots 

from woody branches either below the leaves or in axils of 

older leaves, comprising 3 or 4 flowers subtended by a showy 

involucral bract and (1 -)2 bracteoles; involucral bract subglobose, ovoid, or ellipsoid, consisting of two connate bracts, 

completely enclosing flowers in bud, at anthesis opening along 

one side only and usually remaining subglobose or rarely 

opening wide; staminate pseudanthia sometimes with pistillodes representing reduced pistillate flowers, pistillate pseudanthia sometimes with rudimentary calyx(ces) representing 

reduced staminate flower(s). Staminate flowers sessile; calyx 

present, reduced, or absent, lobed or laciniate; corolla and disc 

absent; stamens (2-)3-4; filaments slender or stout, usually 

connate at base. Pistillate flowers sessile or shortly pedicellate; 

calyx, corolla, and disc absent; ovary 3-locular; styles absent or 

very short and connate; stigmas 3, sessile or subsessile, connate 

into thick peltate structure. Fruit a 3-seeded schizocarpic 

capsule, globose or ellipsoid, dehiscing into 3 woody cocci; 

collumella slender, usually not persistent. Seeds ovoid to 

ellipsoid, radially compressed, carunculate, surface smooth, 

shiny, and dark colored; caruncle usually large and colored, 

usually red. 

DISTRIBUTION.-Approximately 30 species in the Neotropics; three species collected in the Guianas. 

LITERATURE.-P~ and Hoffhann (1 91 9c). 

Pera arborea Mutis.] 
NUMBER 86 

Key to the Species of Peru 

1. Leaf glabrescent, often with scattered simple hairs on petiole and on blade 

undersurface near base; stamens mostly fused into stamina1 column 1.5-2.5 mm 

long; involucral bract subtended by a single bracteole .... I? distichophylla* 

Leaf blade undersurface and petiole with lepidote or stellate hairs; stamens free or 

only partly fused into staminal column less than 1 mm long; involucral bract 

subtended by 2 bractoles ............................. 2 

2. Leaf blade very densely covered with hairs below, blade surface not visible between 

hairs; filaments slender, longer than 2 mm .................... 3 

Leaf blade sparsely to moderately densely covered with hairs below, blade surface 

clearly visible between hairs; filaments stout, shorter than 2 mm ........ 4 

3. Hairs lepidote; leaf blade undersurface and capsule densely covered with flat 

scale-like hairs ........................... 2. I? decipiens 

Hairs stellate; leaf blade undersurface and capsule appearing densely tomentose . . ................................... I? tomentosa** 

4. Capsule glabrous, green or bluish green; ovary glabrous or covered with flat lepidote 

hairs; leaf blade undersurface and petiole sparsely lepidote, hairs distinctly 

scale-like often with an uneven or irregularly fringed margin ... 3. I? glabrata 

Capsule and ovary very densely covered with stellate hairs and appearing tomentose, 

yellowish brown in dried state; leaf blade undersurface and petiole sparsely 

..................................... 1. I? bicolor 

to moderately densely stellate-lepidote, hairs with long radiating arms ...... 

9 

*Although Peru distichophyllu (Mart.) Baill. and f? tomentosu (Benth.) Muell. Arg. have been recorded for 

the Guianas, neither apparently occur there. Both species are found in the neighboring area of Venezuelan 

Guayana, primarily in the state of Amazonas. The record off? distichophyllu in Guyana is based on the 

collection Schomburgk 918, which appears to be mistakenly labelled “British Guiana.” The collection is most 

likely from Amazonas, Venezuela, where Schomburgk also collected extensively. 

**Peru tomentosu originally was based on two discordant types, Schomburgk 114 from Guyana and Spruce 

from “Capoeiras, Barra do Rio Negro” (Amazonas, Brazil). To resolve this problem the taxon recently was 

lectotypified by the Spruce collection, thus excluding Schomburgk 114, the type of f? bicolor (Gillespie, 

1993a). This provides a valid and appropriate name, f? tomentosu, for this distinct Amazonian species having 

a densely stellate-tomentose leaf undersurface. This species has not been collected in the Guianas. 

1. Peru bicolor (Klotzsch) Muell. Arg. 

FIGURE 2A,B,D-I 

Peru bicolor (Klotzsch) Muell. Arg. in DC., Prodr., 15(2):1028, 1866. 

Peridium bicolor Klotzsch, London J. Bot., 2:44, 1843. [Type: Guyana, 

Schomburgk 114 (lectotype, G-DC!, designated by Gillespie, 1993a; 

isotypes, G-2 sheets!, W photo F32519!, K-2 sheets!, P-2 sheets!).] 

Peru schomburgkianu Muell. Arg. in DC., Prodr., 15(2): 1027,1866. [Syntypes: 

GuyanaNenezuela, Roraima, Ri. Schomburgk 901 (not seen), 905 (syntype, 

B, destroyed, photo F5376!); Ro. Schomburgk II 580 (syntype, G-DC!; 

isosyntypes, G-2 sheets!, P-2 sheets!).] 

DESCRIPTION.-Shrub or tree to 33 m, dioecious; indumentum stellate or stellate-lepidote, hairs less than 0.2 mm in 

diameter, with slender radiating arms. Young stems densely 

stellate-lepidote. Petiole 0.3- 1.3 cm long, densely stellatelepidote; blade subcoriaceous, obovate or elliptic, 5-1 1 x 3-6 

cm, shiny dark or bright green above, pale grayish green below, 

drying blackish above and pale tan below, apex rounded or less 

often emarginate, short-acuminate or obtuse, base obtuse or 

acute, glabrous above and often with stellate-lepidote midrib, 

sparsely covered with stellate-lepidote hairs below. Inflorescence peduncle 1.5-4.5 mm long; involucral bract yellow, 

subglobose, 4-6 mm in diameter, densely stellate-lepidote, 

bibracteolate at base. Staminate flowers 3 per involucral bract; 

calyx irregularly laciniate, to 1.3 mm long or less; stamens (3-) 

4; filaments 1-2 mm long, stout, connate at base; anthers -1 

mm long. Pistillate flowers 3 per involucral bract; ovary 

subglobose, indistinctly 3-lobed, -2 mm in diameter, very 

densely tomentose; stigmas obtriangular or obovate, connate at 

base into a deeply 3-lobed structure, 2-3 mm in diameter. 

Infructescence peduncle 0.3-0.4 mm long; pedicel 0.3-0.5 

mm long; capsule ellipsoid-globose, 15-19 x 14-16 mm, 

yellow-brown, very densely stellate appearing densely tomentose. Seeds ovoid, radially compressed, black, -7 x 5-6 x 3 

mm; caruncle red, covering about two-thirds of seed. 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 86 11 

FIGURE 2 (left).-Peru bicolor (Klotzsch) Muell. Arg. and Peru glabrata 

(Schott) Baill. A,B,D-I, P: bicolor: A, habit, flowering branch showing 

fasciculate staminate pseudanthia; B, leaf blade abaxial surface showing 

stellate-lepidote hairs; D, staminate pseudanthium at anthesis, ventral view 

showing slit-like opening of subglobose involucral bract with staminate flowers 

inside; E, staminate pseudanthium at anthesis, side view; F, staminate 

pseudanthium with involucral bract removed showing cluster of three staminate 

flowers; G, staminate flower; H, pistillate pseudanthium with involucral bract 

removed showing cluster of four pistillate flowers; I, cluster of two capsules on 

branch (A,B,D-G based on Gillespie 4300, US'; n based on Huber 5170, US; I 

based on Pinkus 194, US); C, P: glubruru, leaf blade abaxial surface showing 

lepidote hairs (based on Pipoly & Godfrey 7371, US). 

DISTRIBUTION.-Venezuela (Amazonas, Bolivar), the Guianas, and Brazil (Amazonas); 38 collections studied, of which 

26 are from the Guianas (G:20, S:3, FG:3). 

ECOLOGY .-Occurs in savanna, savanndforest edge, or 

seasonal forest (e.g., Wallaba forest in Guyana), on white sand 

or sandy soils, near sea level to 650 m. Flowers have been 

reported from September to November, fruits from February to 

April in the Guianas. Trees have been observed to mass flower; 

small bees were seen visiting and collecting pollen from the 

very fragrant, lemon-scented, yellow pseudanthia of staminate 

trees (Gillespie, pers. obs. in Guyana). 

SELECTED SPECIMENS EXAMINED.4Uyana: Kataima Mazaruni River, Maguire di Fanshawe 32639 (NY, US); mouth of 

Kako River, upper Mazaruni, Pinkus 194 (GH, MO, NY, US); 

Moraballi Creek near Bartica, Sandwith 425 (G, NY, P, US). 

Surinam: Para District, Sectie 0, BW 4785 (A, GH, NY, US); 

Jodensavanne-Mapane Kreek area, Lindeman 4922 (NY). 

French Guiana: La Mana, 1857, Sagot s.n. (G-DC, P); Route de 

Charvein-Acarouany, km 6, Sew. For. 311M (NY). 

VERNACULAR NAMEs.4uyana: Hachiballi (Arawak), Urr 

(Wapisiana). Surinam: Hachiballi, hatsiballi (Arawak), peprehoedoe (Surinamese), pirikraipjo (Carib). French Guiana: 

Baaka tiki (Paramaka). 

NOTES.-There has been much confusion in the literature 

over the identity of P. bicolor, with the name frequently 

misapplied. The confusion appears to begin with Bentham 

(1 854) who grouped three species under Peridium bicolor. In 

his description of P. bicolor var. tomentosum, he cited both the 

type collection of P. bicolor from Guyana and a collection of a 

second species from Brazil, now known as P. tomentosa. A 

third species was treated as variety nitidum (=P decipiens). 

Mueller (1865, 1866) treated P. bicolor (including var. 

nitidum) and P: tomentosa as two distinct species and described 

two additional species, P. decipiens and P. schomburgkiana (= 

P bicolor). Pax and Hoffmann (1919~) treated P: bicolor in the 

sense of variety nitidum (including P. decipiens) and considered both I? schomburgkiana and P. tomentosa as distinct 

species. More recently, in their treatments of Guianan 

Euphorbiaceae, Lanjouw (193 1) and Lemee (1952) treated P: 

bicolor in the correct sense; in addition, Lanjouw noted that 

Peridium bicolor var. tomentosum was based on two discordant 

types (discussed above) and that Peridium bicolor var. nitidum 

belongs under P. decipiens. 

Intraspecific variation in the density of stellate hairs on the 

leaf blade undersurface may be responsible for some of the 

confusion described above. The type specimens of €? schomburgkiana have a sparsely stellate leaf blade undersurface, 

whereas the type specimen of I? bicolor has a more densely 

stellate undersurface. They represent opposite ends of a 

gradation in hair density in this species. 

2. Peru decipiens (Muell. Arg.) Muell. Arg. 

FIGURE 3 

Peru decipiens (Muell. Arg.) Muell. Arg. in DC., Prodr., 15(2):1029, 1866. 

Peridium decipiens Muell. Arg., Linnaea, 34:201, 1865 [based on type of 

Peridium bicolor var. nitida]. 

Peridium bicolor var. nitidum Benth., Hooker's J. Bot. Kew Gard. Misc., 6:323, 

1854. [Type: Guyana. Ri. Schomburgk 1070 (lectotype, K!, designated by 

Gillespie, 1993a; isolectotype, G-fragment!).] 

Peru nitida (Benth.) Jabl., Mem. New York Bot. Gard., 17:148, 1967. 

DESCRIPTION.4hrub or tree to 30 m; dioecious; indumentum lepidote, hairs -0.2 mm in diameter, scale-like with darker 

center and fringed margin. Young stems very densely covered 

with lepidote hairs. Petiole 0.6-1.8 cm long, very densely 

lepidote, with adaxial groove; blade subcoriaceous, narrowly 

elliptic or elliptic, 6-14 x 3-6 cm, dark green above, dull 

yellow or pale green below, apex obtuse, acute, or sometimes 

rounded, base obtuse or acute, glabrous with lepidote midrib 

above, very densely lepidote below. Inflorescence peduncle 

3-8 mm long; involucral bract cream white or pale yellow 

usually becoming pink, globose, 6-8 mm in diameter, densely 

lepidote, bibracteolate at base. Staminate flowers 3 per 

involucral bract; calyx 1-1.5 mm long, irregularly and 

shallowly lobed, sparsely to densely covered with stellate or 

stellate-lepidote hairs, or hairs restricted to margin; stamens 

2-4; filaments 2.2-3 mm long, slender, connate at base; 

anthers 0.7-1 mm long, sagitate. Pistillate flowers 3-4 per 

involucral bract; 1-5 rudimentary staminate calyces often 

present between pistillate flowers; ovary ellipsoid, 2-2.5 x 

-1.5 mm, 3-angled, very densely lepidote; stigmas connate 

into a triangular, peltate structure, 2-3 mm wide. Infructescence peduncle 5-6 mm long, pedicel 5-6 mm long; capsule 

subglobose, 1.0-1.3 cm in diameter, green, very densely 

lepidote. Seeds ovoid, radially compressed, dark brown, -6 x 5 

x 2.5 mm, almost covered by flap-like extension of caruncle. 

DIsTRIBUT1oN.-Venezuela (Amazonas, Bolivar), Guyana, 

Surinam, Brazil (Amazonas, Para, RondBnia); 29 collections 

examined, of which 10 are from the Guianas (G:7, S:3). 

ECOLOGY.4ccurs in seasonal to moist forest, gallery 

forest, savanna, or blackwater inundated forest on sandy soils, 

near sea level to 400 m. Flowering specimens have been 

collected between September and December in the Guianas. 

The creamy white or pale yellow pseudanthia are reported to be 

fragrant. 

SPECIMENS ExAMINED.4uyana: Parabara Savanna, Guppy 

627 = FD 7642 (FDG, NY, P); Kamoa R., Toucan Mt., Jansen 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 86 13 

FIGURE 3 (left).-Peru decipiens (Muell. Arg.) Muell. Arg.: A, vegetative 

branch; B, leaf blade, abaxial surface showing dense covering of lepidote hairs; 

C, flowering branch, showing fasciculate clusters of pistillate pseudanthia; D, 

pistillate pseudanthium at anthesis; E, pistillate pseudanthium with involucral 

bract removed showing cluster of four pistillate flowers; F, staminate flower; G, 

cluster of two dehisced capsules (A,B,F based on Smith 2658, US; C-E based on 

Guppy 267 = FD 7642, MO; G based on Bemy 604, MO). 

Jacobs et al. 1751 (P); “British Guiana,” 1842, Ri. Schomburgk 

1071 (K); Roraima, Ro. Schomburgk 685 (G, K, P), 686 (G, 

G-BOIS, G-DC, K, P); Essequibo River, near mouth of Onoro 

Creek, Smith 2658 (A, NY, P, US). Surinam: margin of Kayser 

Airstrip, Zuid River, 45 km above confluence with Lucie River, 

Irwin et al. 57504 (K, MO, NY, US); N slope of ‘4-Geboiders’ 

Mts., Sipaliwini savanna area, Oldenburger et al. 264 (K, NY); 

N of ‘4-Geboiders’ Mts., Sipaliwini savanna area, Oldenburger 

et al. 318 (NY). 

VERNACULAR NAMES.-Surinam: Peprehoedoe. 

3. Pera glabrata (Schott) Baill. 

FIGURE 2c 

Peru glubrutu (Schott) Baill., Etude Euphorb., 434, 1858. 

Peridium glubrutum Schott in Spreng., Syst. veg., 4(2)(curae post.):410, 1827. 

[Type: none designated (presumably described from one or more specimens 

in Schott’s own collection; Mueller apparently saw these specimens and 

cited them as Schotr (1866: 1030) and later as Schott 4318, 451 7 (1874:430).] 

Peridium ferrugineum Schott in Spreng., Syst. veg., 4(2)(curae post.):4lO, 

1827. [Type: none designated]. 

Peru ferrugineu (Schott) Muell. Arg. in DC., Prodr., 15(2):1031, 1866. 

DESCRIPTION.-Shrub or tree to 37 m, dioecious; indumentum lepidote, hairs 0.2-0.4 mm in diameter, scale-like with 

uneven and sometimes irregularly dissected or fringed margin. 

Young stems sparsely to moderately lepidote. Petiole 0.5-1.2 

cm long, sparsely to moderately lepidote; blade chartaceous to 

subcoriaceous, elliptic, 5-12 x 2.5-5.5 cm, shiny dark green 

above, dull pale green below, apex obtuse, broadly acute or 

broadly acuminate, base obtuse or rounded, glabrous above 

sometimes with sparsely lepidote midrib base, sparsely 

lepidote below. Inflorescence peduncle 3-7 mm long; involucral bract pale yellow or creamy white, ellipsoid or subglobose, 

3-6 mm in diameter, lepidote, bibracteolate at base. Staminate 

flowers 3 per involucral bract; calyx absent; stamens (3-)4; 

filaments 0.5-1 mm long, stout, connate at base; anthers 1-1.5 

mm long. Pistillate flowers (3-)4 per involucral bract; ovary 

ellipsoid, -1.5 x 1 mm, covered with lepidote and sometimes 

stellate-lepidote hairs or rarely glabrous; stigmas connate into 

a 3-lobed peltate structure, -1 mm wide, distinctly vermcosepapillate. Infmctescence peduncle 6- 12 mm long, pedicel 3-5 

mm long; capsule broadly ellipsoid, 8-13 x 7-11 mm, 

blue-green or pale green, glabrous, smooth, becoming 

reticulate-wrinkled when mature. Seeds ovoid, radially compressed, black, 4-6 x 3-4.5 x 2-3.5 mm, apex distinctly 

narrowed; caruncle orange or red, mostly covering adaxial seed 

surface. 

DISTRIBUTION.-Venezuela, the Guianas, Brazil (Amazonas, Para, eastern and southeastern Brazil); 63 collections 

studied, of which 25 are from the Guianas (G: 17, S:2, FG:6). 

ECOLOGY.-Occurs in seasonal lowland to montane forest, 

often on white sand, sea level to 700 m in the Guianas, to 1400 

m in Venezuela. Flowers have been reported in March, April, 

September, and November, fruit in May and November in the 

Guianas. The pale yellow or creamy white pseudanthia are 

reported to be fragrant. 

SELECTED SPECIMENS ExAMINED.4uyana: Mazaruni Station, FD 5409 (K, US); Mainstay Village, 7’15’N, 58”32’W, 

Gillespie & Persaud 1058 (US); Linden-Soesdyke Hwy., 

between Dora and Maibia Creek, 6’18’N, 58”14’W, Pipoly et 

al. 9705 (NY, US); Basin of Rupununi R., Isherton, Smith 2418 

(G, US). Surinam: Kaboerie, BW 4847 (A, GH, K, US); 

Zanderij I., Stahel273 (A, GH, K). French Guiana: Commune 

de Iracoubo, 5”28’N, 53”14’W, Cremers 9627 (US); 

Ile de Cayenne, route de Rochambeau, Oldeman BC9 (MO, 

VERNACULAR NAMES.4uyana: Hatchi-balli (Arawak), 

NY, US). 

kamua-tan (Wapisiana). Surinam: hatsiballi koelnoe. 

Tribe PLUKENETIEAE 

Tribe Plukenetieae (Benth.) Hutch., is a cohesive group of 14 

genera distributed worldwide in tropical and warm temperate 

regions. The tribe may be distinguished by its bisexual, 

racemose or spicate inflorescences (unisexual in Acidoton Sw. 

and some African Tragia spp.), and entire styles that are basally 

to completely connate and often massive (most other members 

of the Acalyphoideae core group have unisexual inflorescences 

and styles that are typically branched and/or free). In addition, 

many species have a scandent habit and/or urticating hairs, both 

unusual features in the Euphorbiaceae. 

Tribe Plukenetieae comprises three subtribes, the Dalechampiinae, Plukenetiinae, and Tragiinae (Webster, 1994). The 

Tragiinae have urticating hairs, a trilocular ovary, and lack 

laminar glands, whereas the Plukenetiinae lack urticating hairs 

and have a 3- or 4-locular ovary and usually laminar glands on 

the leaf blade. Pollen of the Plukenetiinae is uniformly 

tricolpate, whereas that of the Tragiinae varies from tricolpate 

to weakly aperturate and inaperturate (Gillespie, 1994a). 

Tragiinae genera known from the Guayana Region include 

Acidoton and Tragia, whereas genera of Plukenetiinae known 

from this region include Astrococcus Benth., Haematostemon, 

and Plukenetia (Gillespie, 1993a). Of these genera, only 

Haematostemon, Plukenetia, and Tragia have been collected 

from the Guianas. Acidoton may perhaps be expected to occur 

in the Guianas, having been collected from the neighboring 

state of Bolivar, Venezuela, and is therefore included in the 

key. Astrococcus, on the other hand, is endemic to blackwater 

inundated forest in the upper Rio Negro region and is unlikely 

to be found in the Guianas. 

The third subtribe, Dalechampiinae, comprising the single 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

genus Dalechampia, is unique in the Euphorbiaceae in having 

2-bracted, bisexual pseudanthia. The subtribe, previously 

considered a distinct but related tribe (Webster, 1975), has 

recently been placed in tribe Plukenetieae (Webster, 1994). The 

presence of urticating hairs, styles that are completely connate 

into a thickened, often massive column, bisexual inflorescences, and a usually scandent habit strongly suggest a close 

affinity with other members of tribe Plukenetieae. 

3. Genus Dalechampia L. 

Dalechampia L. Sp. PI., 1054, 1753; Gen. PI., ed. 5:473, 1754. [Type: 

Megdosfylis Spencer Moore, J. Bot., 54:208, 1916. [Type: Megalosfylis 

DESCRIPTION.-Perennial vines or shrubs, monoecious; 

latex absent. Stems climbing, spreading, or erect, usually 

covered with urticating hairs. Leaves simple or compound, 

alternate, petiolate, stipulate, with pair of stipels at base of 

blade (stipels sometimes obscure or deciduous); blade pinnately or palmately veined, margin entire to deeply lobed; 

distinct petiolar and laminar glands absent. Inflorescence 

pseudanthial, subtended by 2 usually large, showy bracts, 

Dalechampia scandens L.] 

poeppigii Spencer Moore.] 

comprising a cymule of 3 pistillate flowers and 3-5-branched 

pleiochasium of 4-(-15) staminate flowers; pistillate cymule 

subtended by an involucel of 1 bract and 0-2 bractlets; 

staminate pleiochasium subtended by a cup-like involucellar 

bract, a bilabiate involucel, or 4(-5) free bracts. Staminate 

flowers with calyx splitting into 3-6 valvate segments at 

anthesis; corolla and disc absent; stamens 5-90 on flat, 

dome-shaped, or elongated receptacle (column); anthers subsessile or on short filaments, usually in cluster at apex of 

column, dehiscing longitudinally. Pistillate flowers with 5- 12 

entire or lobed sepals; petals and disc absent; ovary 3-locular; 

styles connate into elongated column, often dilated at apex. 

Fruit 3-seeded capsule, dehiscing explosively by elastic 

twisting of dry, more or less woody cocci. Seeds subglobose to 

globose, ecarunculate, surface smooth, roughened, or tuberculate, usually gray-brown mottled. 

DISTRIBUTION.-Approximately 120 species, -95 in neotropics, 10 in Africa, 10 in Madagascar, and six in Asia; 16 species 

collected in the Guianas, with two additional species from 

neighboring Venezuelan Guayana likely to occur in the 

Guianas. 

LITERATURE.-Pax and Hoffmann (1 9 19b); Webster and 

Armbruster (1991). 

Key to the Species of Dalechampia 

1. Shrubs; leaves pinnately veined; involucral bracts pink (rarely yellowish) at 

anthesis; inflorescence glands yellow or greenish, surface papillate, producing 

spicy fragrance ........................ 8. D. magnoliifolia 

Vines; leaves palmately veined; involucral bracts white, green, or pink at anthesis; 

inflorescence glands, when present, variously colored, usually producing resin 

........................................... 2 

2. Leaves compound, 3-foliolate or rarely 4- or 5-foliolate; involucral bracts 

conspicuous, green at anthesis .......................... 3 

Leaves simple; involucral bracts inconspicuous or conspicuous, white, pink, or 

green at anthesis ................................. 5 

Involucral bracts spathulate, 5-8 mm long, 3-4 mm wide, deciduous prior to 

fruiting; staminate involucellar bracts free, 4 in number; surface of stigmas 

papillate ............................ 12. D. papillistigma 

Involucral bracts broadly ovate, 10-15 mm long, 10-15 mm wide, persistent in 

fruit; staminate involucellar bracts connate or bilabiate; surface of stigmas smooth 

........................................... 4 

Pistillate sepals shallowly to deeply lobed, rachis broad, width >l/4 length, fringed 

with dense pubescence .................... 18. D. aff. triphylla 

Pistillate sepals pinnatifid, rachis narrow, width <l/4 length, hispid ........ 

5. Leaves unlobed; involucral bracts subequal, inconspicuous, small (C6 mm long), 

stipuliform, green at anthesis, or dimorphic (one small, the other conspicuous, >1 

cm long, whitish); inflorescence gland usually absent; pistillate sepals 6, unlobed 

........................................... 6 

Leaves lobed or unlobed; involucral bracts subequal, conspicuous, 21 cm in length; 

resin-producing inflorescence gland present (vestigial and inconspicuous in D. 

brownsbergensis); pistillate sepals 6-12, lobed or unlobed .......... 12 

3. 

4. 

................................. 17. D. aff. cissifolia 
NUMBER 86 15 

6. Leaves narrow, width 413 length, with arcuate secondary veins branching from 

basal and distal regions of the central vein, base of blade obtuse and rounded; 

staminal column extremely short or obsolete; stylar column cylindrical, slightly 

dilated at extreme tip ..................... 3. D. brevicolumna 

Leaves wide, width 9/3 length, with 1-3 arcuate secondary veins attached to the 

central vein only in its distal '12, base of blade truncate to cordate; staminal column 

well developed, conspicuous; stylar column enlarged, more or less clavate, not 

dilated at extreme tip ............................... 7 

Stems, petioles, leaves, and peduncles densely hirsute-lanate with orange hairs; 

pseudanthia subsessile, usually 2 or more in leaf axils ..... 11. D. olympiana 

Stems, petioles, leaves, and peduncles glabrescent to hirsute, but hairs not orange; 

pseudanthia borne on peduncles (short shoots) >1 cm long, usually solitary in leaf 

axils ........................................ 8 

8. Base of leaf blades truncate to shallowly cordate; stipules ovate, 5 mm long, early 

deciduous; fertile arms in staminate pleiochasium 4; staminate flowers 12- 13; 

Base of leaf blades shallowly to deeply cordate (or hastate in immature plants); 

stipules lance-ovate to lanceolate, usually >5 mm long (except D. purvibructeutu), 

deciduous only after leaves have expanded, or persistent; fertile arms in staminate 

pleiochasium 3-4; staminate flowers 10-13; stigma extends from tip to <3/4 

length of style or if to >3/4 length, then widest part of stylar column C1/2 distance 

from base; pistillate sepals persistent into fruit ................. 10 

Stylar column curved upwards toward staminate cymule, width -l/5 length, tip 

markedly foveolate; base of leaves usually truncate, stipules deciduous prior to 

full expansion of leaves ..................... 10. D. micrantha 

Stylar column more or less straight, very thick, width -'/3 length, tip not markedly 

foveolate; base of leaves usually shallowly cordate, stipules deciduous usually 

after expansion of leaves ...................... 6. D. fragrans 

10. Stylar column broadest at '/3 to '12 length from base, tapering to blunt, attenuate tip; 

smooth stigmatic surface extending from tip to 2/3 to 3/4 length of stylar column, 

not contrasting markedly with stylar base; sinus at base of leaves usually deltoid, 

narrow at attachment of blade to petiole ........... .2. D. attenuistylus 

Stylar column clavate, broadest at w2/3 to 3/4 its length from base; smooth stigmatic 

surface extending from tip to l/4 to 2/3 length of stylar column, contrasting 

markedly with pubescent stylar base; sinus at base of leaves usually broad and 

U-shaped ..................................... 11 

11. Involucral bracts usually dimorphic, the one subtending pistillate flowers >1 cm 

long, whitish, the one subtending staminate flower <5 mm long, stipuliform, 

green; stigmatic surface extending from tip to w2/3 length of stylar column 

................................. 7. D. heterobractea 

Involucral bracts subequal, <5 mm long, stipuliform, green; stigmatic surface 

extending from tip to -'I4 length of stylq column ..... 13. D. parvibracteata 

12. Leaves always unlobed .............................. 13 

Leaves all 3-lobed or both unlobed and 3-lobed present on same plant (except very 

young plants, which may have only unlobed leaves) .............. 16 

Staminate pleiochasium with 4 free involucellar bracts; inflorescence gland 

comprising concentrically arranged bractlets with laciniate margins, secreting 

greenish or brownish resin; involucral bracts linear and green or ovate and pink 

.......................................... 14 

Staminate pleiochasium with involucel of bracts fused for at least basal l/3 or 

forming a bilabiate cup; inflorescence gland of more or less laminar bractlets with 

entire or minutely fimbriate margins, secreting clearish or yellowish resin; 

involucral bracts broadly ovate, pale green or white .............. 15 

7. 

stigma extending from tip to >3/4 length of style ................ .9 

9. 

13. 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

14. Leaf blades 7-27 cm long with brownish pubescence below, involucral bracts green 

at anthesis, 1-2 cm long, inflorescence gland secreting bluish green resin, style 

more or less evenly columnar, pistillate sepals 6, unlobed, fruit 3.5-4 cm in 

diameter. ............................ 9. D. megacarpa 

Leaf blades 5-15 cm long, lacking brownish pubescence, involucral bracts pale 

pink with reddish veins, 3-5 cm long, inflorescence gland secreting brownish 

maroon resin, style with distinct peltate stigmatic tip, pistillate sepals 6- 12, 

twice-pinnatifid, fruit C2 cm in diameter. .......... 5. D. dioscoreifolia 

15. Involucral bracts white at anthesis, >3 cm long, entire or shallowly 3-lobed, with 

7-9 prominent veins; inflorescence gland secreting yellowish resin; tips of styles 

broadly dilated and symmetrically peltate .............. 1. D. affinis 

Involucral bracts pale green to whitish at anthesis, c3.5 cm long, deeply 3-lobed to 

-*/2 their length, with 5 prominent veins; inflorescence gland secreting clear to 

whitish resin; tips of styles modestly and asymmetrically dilated ......... .......................... 14a. D. scandens var. scandens 

16. Margins of leaf stipules, involucral bracts, and (usually) leaf blades capitate 

glandular, glandular appendages secreting sticky resin ..... 15. D. stipulacea 

Margins of leaf stipules, involucral bracts, and leaf blades subentire to irregularly 

17. Leaf stipules ovate to lance-ovate, persistent; involucral bracts 3-lobed to -*/2 their 

length, C3.5 cm long at anthesis, with 5 prominent veins; inflorescence gland 

secreting clear to whitish resin ......................... 18 

Leaf stipules lance-ovate to lance-linear, deciduous; involucral bracts shallowly 

3-dentate at apex, >3 cm long at anthesis, with 7-9 prominent veins; inflorescence 

18. Young stems and petioles with sparse to moderately dense spreading straw-colored 

pubescence; underside of mature leaves glabrescent to strigulose; involucral bracts 

pale green to whitish, margins with small fimbria, these at most weakly capitate, 

c0.5 mm long; margin of proximal bract of pistillate involucel entire; stylar 

.......................... 14a. D. scandens var. scandens 

Young stems and petioles with very dense spreading orange pubescence; underside 

of mature leaves lanate; involucral bracts white, margins with strongly capitate 

tentacular processes, generally 1-1.5 mm long; margins of proximal bract of 

pistillate involucel capitate glandular; stylar column with broadly expanded 

19. Leaf stipules lance-linear; involucral bract whitish cream at anthesis; inflorescence 

Leaf stipules lance-ovate to lanceolate; involucral bracts pale green at anthesis; 

............................... 4. D. brownsbergensis 

dentate, but without glandular appendages that secrete sticky resin ...... 17 

gland inconspicuous or secreting yellowish resin ................ 19 

column with slightly and asymmetrically dilated tip ................ 

symmetrically peltate tip .............. 14b. D. scandens var. fallax 

gland prominent, secreting yellowish resin ............ 16. D. tiliifolia 

inflorescence gland inconspicuous and vestigial, not secreting resin ....... 

1. Dalechampia affinis Muell. Arg. 

Dalechampia aflnis Muell. Arg., Linnaea, 34:223, 1865. [Type: French 

Guiana, La Mana, Sagor s.n. (lectotype, G, designated by Webster and 

Armbruster, 1991).] 

DESCRIPTION.-Twining or sprawling vine. Leaves simple; 

stipules lance-linear, 3-7 x -1 mm; petioles 2-6 cm long; 

blades chartaceous, usually ovate, rarely lanceolate, always 

unlobed, 4-12 x 3-9 cm, apex acuminate, base shallowly to 

white at anthesis, broadly ovate, tip acuminate, usually entire, 

2-5 x 2-4 cm, velutinous, with 7-9 major veins; pseudanthium containing -10 staminate flowers and 3 pistillate 

flowers; inflorescence gland secreting yellow resin. Staminate 

flowers with -40 stamens. Pistillate flowers with calyx of 

10- 12 sepals, these pinnatifid, densely hispid; pistil with 

elongate stylar column, 6-10 mm long, tip symmetrically 

umbraculiform. Fruit surface sparsely to densely hispid; seeds 

spherical, mottled brown and gray, -4 mm in diameter, surface -. 

deeply cordate, sparsely pubescent above, velutinous below. 

Inflorescence pseudanthial; the two bracts involucral, creamsmooth. 

DISTRIBUTION.-Widespread in lowland northeastern South 
NUMBER 86 17 

America (eastern Venezuela, Guyana, Surinam, French Guiana, 

and northern Brazil); 36 collections studied, all from the 

Guianas (G:10, S:13, FG:13). 

ECOLOGY.4ccurs in open woodland and savanna below 

500 m. The copious yellow resin secreted by the inflorescence 

glands is collected in Brazil by female euglossine bees 

(including Euglossa; Apidae) that pollinate flowers in the 

process (Armbruster and Webster, 198 1, 1982). It apparently 

flowers December through July. 

SELECTED SPECIMENS ExAMINED.aUyana: Naamryck 

Canal, Essequibo Islands-West Demerara, -3.5 km SW of 

Parika, 6"50'N, 58"27'W, Gillespie & GopaullOO9 (ALA, US); 

Rockstone, bank of Essequibo River, Gleason 880 (NY, US). 

Surinam: Nickerie, near Kabalebo Dam project, Lindeman et 

al. 286 (K, MO, NY, U, US); Zuid River, between Kayser 

airstrip and confluence with Lucie River, 3"20'N, 56'49'W3"10'N, 56"29'W, Maguire et al. 54028 (MO, NY, U, US). 

French Guiana: Village Eskol, 4"45'N, 52"20'W, Cremers 9820 

(CAY, NY, US); Tampoc River, SE of Maripasoula, Moretti 

698 (CAY). 

NOTES.-This species is similar and very closely related to 

D. tiliifolia. From D. tiliifolia it differs primarily in having 

exclusively unlobed leaves, symmetrically peltate style tips, 

and involucral bracts usually with unlobed tips. 

2. Dalechampia attenuistylus Armbruster 

Dalechampia attenuistylus Armbruster, Brittonia, 41 :44, 1989. [Type: Venezuela, Bolivar, Santa Elena de Uairen, Armbruster et al. 87-153 (holotype, 

MO!; isotypes, ALA!, MY!, NY!, VEN!).] 

DESCRIPTION.-Twining vine reaching low forest canopy. 

Leaves simple; stipules lanceolate to ovate, 3-8 x 1-2 mm; 

petioles 1-4.5 cm long; blades unlobed, 3-10 x 2-7 cm, base 

cordate with deltoid sinus, sinus usually narrow at attachment 

of blade to petiole. Pseudanthial inflorescences with inconspicuous stipuliform involucral bracts 3-6 mm long, rarely 

lower (basal) bract up to 1 cm long, lo(-1 1) staminate flowers 

and 3 pistillate flowers per infloresecence. Staminate flower 

with 8-lo(-1 1) stamens. Pistillate flowers with 6 sepals, these 

unlobed; stylar column straight, widest at lh-lh length from 

base, distally tapered to attenuate, blunt tip; stigma light green, 

extending from tip to 2/3-3/4 length of stylar column. Fruit with 

small persistent sepals; seeds subspherical, -4 mm in diameter, 

surface nearly smooth. 

DISTFUBUTION.-hOWn only from southeastern Bolivar, 

Venezuela, and southern Surinam, but likely to occur in 

Guyana and northern Brazil; three collections studied, one from 

Surinam. 

ECOLOGY.-Vine in forest openings and lower canopy of 

moist riparian forest, 800-1000 m. In flower and fruit middle 

to late December. 

SPECIMENS EXAMINED.-sUIinam: Lely Mountains, Mori 

and Bolten 8550 (MO, NY, U, US). Venezuela: Bolivar, Santa 

Elena de Juairen, first bridge south of town, Armbruster et al. 

NOTES.-A member of section Rhopalostylis, this species is 

probably most closely related to D. pawibracteata. As is the 

case for all members of the section, it lacks the prominent 

inflorescence gland characteristic of most of the rest of genus. 

It differs from other members of the section in having tapered 

styles that are widest at about one-third the length and in having 

10 staminate flowers arranged in 3 pleiochasial arms in the 

inflorescence. 

93-01 (ALA). 

3. Dalechampia brevicolumna Arm b rus ter 

FIGURE 4 

Dalechampia brevicolumna Armbruster, Syst. Bot., 2 1:232, 1996. [Type: Near 

St. Elie, French Guiana, Prevost 661 (holotype, CAY!; isotype, U!).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules lanceolate, 2-4 x 0.8-1 mm; petioles 4-20 

mm long; blades 5-11 x 2-5 cm, width C l/3 length, base 

rounded, triplinerved, secondary veins diverge from central 

primary vein throughout its length. Pseudanthial inflorescences 

with inconspicuous stipuliform involucral bracts 2.5-3.0 mm 

long, 6-9 staminate flowers, 3 pistillate flowers. Staminate 

flowers with 20-30 stamens attached to more or less flat 

receptacle, column extrememly short or absent. Pistillate 

flowers with 5-6 sepals, these lanceolate, unlobed; stylar 

column more or less cylindrical, straight, with slightly dilated 

tip, stigma extending from tip to > 3/4 length of stylar column. 

DIsTF3BuTIoN.-Apparently endemic to French Guiana and 

Surinam; five collections studied (FG:4, S: 1). 

ECOLOGY.4ccurs in secondary vegetation and at edge of 

wet forest. In French Guiana it is pollinated by female 

resin-collecting anthidiine bees (Hypanthidium sp., Megachilidae) and conspecific males landing on the flowers in search of 

females. This species appears to have peak flowering in late 

June through August, although flowers also have been 

collected in March; fruits are present June to October. 

SPECIMENS EXAMINED.-French Guiana: Saul-Belizon area, 

Oldeman 1472 (CAY); near St. Elie, Prevost 665 (CAY, U); 

near St. Elie, Prevost 2673 (US). Surinam: Tumuc Hamuc 

Mountains, Talouaken, Acevedo-Rodriguez et al. 6026 (US). 

NOTES.-This is a very unusual species. It is placed in its 

own section, Brevicolumnae (Armbruster, in press). It resembles members of sect Rhopalostylis in having small, stipuliform 

involucral bracts and leaf lamina with sparse pubescence. It 

differs from members of the section in its leaves having 

secondary veins distributed along the length of the central 

primary vein, the arrangement of staminate flowers and 

involucellar bractlets, and staminate flowers with extremely 

short stamina1 columns. It resembles more advanced species in 

section Dalechampia (including D. scandens and D. stipulacea) in having inflorescence resin glands comprising laminar 

bractlets with entire margins. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 4.-Dulechumpiu brevicolumnu Armbruster: A, habit; B, habit with infructescence in post-dehiscence 

stage; C. inflorescence, pistillate side; D, inflorescence with resin gland, staminate side with part of involucellar 

bract removed; E, pistillate flower; F, staminate pleiochasium lacking resin gland, as is occasionally observed; G, 

staminate flower (based on Prevost 661). 
NUMBER 86 19 

4. Dalechampia brownsbergensis Webster 

& Armbruster 

FIGURE 5

Dalechampia brownsbergensis Webster & Armbruster, Syst. Bot., 7:484, 1982. 

[Type: Surinam, Brokopondo, Brownsberg Nature Preserve, 4"57'N, 

55'1 l'W, Webster & Armbruster 24124 (holotype, DAV!; isotypes, ALA!, 

BBS!, GH!, MO!, NY!, U!). 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules elliptic-lanceolate, 4.5-10 x -2-3.5 mm; 

petioles 4.5-13 cm long; blades chartaceous, ovate, usually 

3-lobed, 10-22 x 10-25 cm, apex of middle lobe cuspidate, 

base deeply cordate, glabrescent above, puberulent below. 

Inflorescence pseudanthial; the two bracts involucral, pale 

green at anthesis, broadly ovate, tip shallowly 3-lobed, 2-5 x 

2-5 cm, 9-11 veined at base, velutinous; resiniferous gland 

inconspicous, vestigial, not secreting resin. Staminate flowers 

with 5-22 stamens. Pistillate flowers with calyx of 10-12 

sepals, these pinnatifid, densely hispid; pistil with elongate 

stylar column, 7-10 mm long, tip asymmetrically trigonous. 

Fruit a capsule, surface glabrate, prominently raised-reticulate; 

seeds spherical, mottled brown and gray, -3.7 mm in diameter, 

surface smooth. 

DISTRIBUTION.-oCCUrS in Surinam and eastern Venezuela, 

also probably Guyana. Five collections studied, two of which 

are from Surinam. 

ECOLOGY.4ccurs in wet forests below 100 m. This species 

lacks the resiniferous inflorescence gland characteristic of most 

species. It instead secretes a fragrance from the stigmatic 

surface of the pistillate flowers and attracts pollinating male 

euglossine bees that collect the fragrance (Armbruster et al., 

1992). Apparently flowers year-round; flowers and fruits are 

present in July, November, and December. 

SPECIMENS EXAMINED.-Surinam: Brokopondo, Brownsberg Nature Park, -4"55'N, 55OlO'W, Armbruster & Herzig 

85- 1 02 (ALA) . 

5. Dalechampia dioscoreifolia Poep p. 

FIGURE 6 

Dalechampia dioscoreifolia Poepp., Nov. gen. spec. pl., 3:20, 1841. [Type: 

Peru, Maynas, Poeppig 2163 (holotype, B, destroyed; lectotype, W, 

designated by Webster and Armbruster, 1991).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules lanceolate, 3.0-3.5 x 1.0-1.5 mm wide; 

petioles 2-7 cm long; blades chartaceous, ovate, unlobed, 

5.5-13.5 x 4.0-9.5 cm, base distinctly cordate, tip acuminate. 

Pseudanthial inflorescence with showy pink involucral bracts, 

these ovate, 1.3-4.5 x 1.2- 3.0 cm, with 3-5 major veins, base 

clawed, usually 10 staminate flowers and 3 pistillate flowers; 

inflorescence gland secretes brown-maroon resin. Staminate 

flowers with 20-30 stamens. Pistillate flowers with umbraculiform stylar column, 3-6 mm long, expanded tip symmetrically 

peltate-disciform; stigmatic surface confined to the expanded 

stylar tip. Seeds subspherical and weakly 3-angled, -5 mm 

long. 

DIsTRIBUTION.-occurs from Nicaragua to Peru; 16 collections studied, eight of which are from French Guiana. 

ECOLOGY.-This robust vine grows at the edge of forest or 

into the canopy of moist to wet primary or secondary forest 

below 1200 m. In Mesoamerica and Peru it is pollinated by 

female Eulaema spp. and Euglossa spp. that collect resin from 

the inflorescence gland (Armbruster and Herzig, 1984). 

Flowers present from January through August. 

SPECIMENS EXAMINED.-French Guiana: Saul, Beekman 

59 (CAY); Saul, route to Belizon, Feuillet 417 (CAY); 

Grand Inini River, east of Maripasoula, Granville 7292 (B, 

CAY, MO, P); Saul, Maas et al. 2226 (CAY, U); Crique 

Ket-oko, NE of Saul, Oldeman 2067 (CAY, MO, NY, P); Saul 

area, Oldeman B.3227 (CAY, NY, U); Saul, Raynal-Rogues 

20006 (CAY). 

6. Dalechampia fragrans Armbruster 

Dalechampiafiagrans Armbruster, Bot. J. Linn. SOC. London, 105: 150, 1991. 

[Type: Surinam, Brokopondo, Brownsberg Nature Park, 4"55'N, 55O 1 O'W, 

Armbruster & Herzig 85-101 (holotype, MO!; isotypes, ALA!, NY!, 

UVS!).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules ovate, 3-4 x 1-2 mm, usually deciduous prior 

to full expansion of leaves; petiole 1-5 cm long; blade ovate, 

unlobed, 6-14 x 4-12 cm, with shallowly cordate base, usually 

with broad U-shaped sinus. Pseudanthial inflorescences with 

inconspicuous stipuliform involucral bracts < 5 mm long, 

usually 13 staminate flowers, and 3 pistillate flowers. Staminate flowers with 4-5 stamens. Pistillate sepals 6, unlobed; 

stylar column clavate, straight, widest w2/3 of the way to the 

rounded tip, 7-10 mm long, stigma green, extending from tip 

to w3/4 length of stylar column. 

DISTRIBUTION.-Known from Surinam, French Guiana, and 

Amapa, Brazil; five collections studied (S:3; FG: 1). 

ECOLOGY.4ccurs in canopy of, or gaps in, lowland wet 

forest. Flowers in early to middle November. Members of this 

species secrete a fragrance from the stigmatic surface of the 

pistillate flowers, attracting (in Surimane) a number of species 

of male euglossine bees (Hymenoptera: Apidae), including 

Eulaema spp., Eufriesea spp., and Euglossa spp. These 

fragrance-collecting bees pollinate the flowers (Armbruster et 

al., 1992). 

SPECIMENS EXAMINED.-Surinam: Brokopondo, Brownsberg Nature Park, 4"55'N, 55"10'W, Armbruster and Herzig 

85-104 (ALA); Brownsberg, Maas et al. 2320 (MO). 

French Guiana: -52 km south of Cayenne, Mori et al. 8693 

(Ny, PI. 

NOTES.-This species is a member of section Rhopalosylis 

and lacks the prominent resiniferous inflorescence gland 

characteristic of most members of the genus. 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 5.-Dalechampia brownsbergensis Webster & Armbruster: A, habit with inhctescence; B, inflorescence 

(based on McDowell4318). 
NUMBER 86 21 

FIGURE 6.-Dalechampia dioscoreijblia Poepp.: A, habit; 8, inflorescence; C, staminate flower; D, capsule, note 

persistent style and fimbriate calyx (A,D based on Williams 303; B based on Armbruster & Henig 79-212; c, 

based on Bell et al. 93-326). 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 86 23 

FIGURE 7 (lefi).-Dalechampia parvibracteata Lanj. and Dalechampia 

heterobractea Armbruster. A,B, D. parvibracteata: A, habit; B, pistillate 

flower (A,B based on McDowell 2264); C-F, D. heterobractea: C, habit; D, 

inflorescence; E, pistillate flower; F, staminate flower (with all but two 

anthers removed) (C based on Feuillet 3015; D-F based on Armbruster et al. 

8 7- I5 7). 

7. Dalechampia heterobractea Armbruster 

FIGURE 7C-F 

Dalechampia heterobractea Armbruster, Syst. Bot., 2 1 :226, 1996. [Type: 

Guyana, Mazaruni Station, by Karau Creek, Sandwith 1565 (holotype, K!; 

isotypes, BM!, NY!, P!, U!, US!).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules lanceolate, (3)5-7 x 1-1.5 mm, persistent; 

petioles 3-9 cm long; blades 6-19 x 5-15 cm, generally 

broadest at '14 to l/2 distance form base, base deeply cordate, 

with broad U-shaped sinus, usually wider than deep (base of 

blade hastate in young plants). Pseudanthial inflorescences 

with dimorphic involucral bracts, upper (distal) inconspicuous, 

green, and stipuliform, 2.5-5 x 1-2 mm, lower showy, whitish, 

10-16 x 5-6 mm; staminate flowers usually 10; pistillate 

flowers 3. Staminate flowers with 10- 1 1 stamens. Pistillate 

flowers with 6 unlobed sepals, sepals obovate-oblanceolate, 

3-4 mm long; stylar column, clavate, straight, widest -l/2 to 2/3 

of the way to the rounded tip, 4-9 mm long; stigma whitish, 

extending from tip to -2/3 length of stylar column. Seeds 

subspherical, 3.5-4.1 mm in diameter. 

DISrRIBUTIoN.-~enezuela, Guyana, Surinam, French Guiana, and northern Brazil; 33 collections studied, 17 ofwhich are 

from the Guianas (G:4, S:7, FG:6). 

ECOLOGY.-In wet forest below 500 m. Found in forest 

margins and young secondary vegetation. Flowers and fruit 

collected year-round. 

SELECTED SPECIMENS EXAMINED.+Uyana: Mazaruni Station, FD 3923 (NY). Surinam: Marowijne, near Moengo, 

Lindeman 6129 (K, NY, U); Oelemari River, 3"6'N, 54"33'W, 

Wessels Boer 958 (U). French Guiana: Tampoc River, -50 km 

SW of Saul, Granville B.4846 (CAY, P, U, US); Village 

NaNouk, Litani and Maroni rivers, Moretti 572 (CAY, U). 

NOTES.-A member of sect. Rhopalostylis, this species lacks 

the prominent resiniferous inflorescence gland characteristic of 

the genus. Members of this common species previously have 

been assigned to D. pawibracteata, but they differ from the 

type of that species in having dimorphic involucral bracts and 

a whitish stigmatic surface that extends further down the style. 

8. Dalechampia magnoliifolia Muell. Arg. 

Dalechampia magnoliifolia Muell. Arg., Linnaea, 34:219, 1865. [Type: Brazil, 

Para, Martius (holotype, B, destroyed; lectotype, M, designated by Webster 

& Armbruster, 1991).] 

Dalechampia roezliana var. amazonica Ule, Verh. Bot. Vereins Prov. 

Brandenburg, 50:82, 1909, 

Dalechampia spathulata var. amazonica (Ule) Pax & K. Hoffm., Pflanzenr., 

IV.147.XI1, 68:11, 1919. 

Dalechampia amazonica (Ule) Macbride, Candollea, 8:26, 1940. [Type: Brazil, 

Amazonas, Jurua Miry, Ule 5865 (lectotype, G, designated by Webster & 

Armbruster, 1991).] 

DESCRIPTION.-Monopodial subshrub 0.5-1.2 m tall. 

Leaves simple; stipules lance-ovate, 3-8 x 2-4 mm; petioles 

0.2-1 .O cm long; blades chartaceous, narrowly oblong to 

oblanceolate, unlobed, 12-30 x 3.5-9.0 cm, base cordate. 

Pseudanthial inflorescence with showy pink involucral bracts, 

these ovate, 2.5-3.5 cm x 1.5-4.0 cm, with 3 major veins, base 

weakly clawed or truncate; usually 10 staminate flowers and 3 

pistillate flowers; inflorescence gland bright yellow, dry, 

secreting spicy fragrance. Staminate flowers with 12- 18 

stamens. Pistillate flowers with columnar style, 4-8 mm long, 

tip very slightly dilated; stigmatic surface extending from the 

tip down most of the length of the stylar column. Seeds 

tuberculate, 6.5-7.0 mm long. 

DISTRIBUTION.-In scattered sites in Amazonian Peru and 

Brazil, eastern Venezuela, and French Guiana; seven collec - 

tions studied, three of which are from French Guiana. 

ECOLOGY.-This diminutive subshrub occurs on floor of 

wet primary or secondary forest, below 250 m. The inflorescence gland secretes fragrance rather than resin. In Peru the 

flowers are pollinated by fragrance-collecting male euglossine 

bees (Eulaema and Euglossa; Armbruster et al., 1989). It 

probably flowers year-round. 

SPECIMENS EXAMINED.-French Guiana: -70 km SSW of 

Camopi, Granville 317 (CAY); confluence of Litani and 

Marouini rivers, Cremers 4998 (CAY, U); -50 km W of 

Oiapoque, Cremers 7075 (CAY). 

9. Dalechampia megacarpa Armbruster 

Dalechampia megacarpa Armbruster, Brittonia, 41:47, 1989. [Type: Venezuela, Bolivar, San Ignacio, Armbruster et al. 85-115 (holotype, MO!; 

isotypes, ALA!, MY!, NY!, VEN!).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules lanceolate to ovate, 2.5-5 x 1.2-1.7 mm wide, 

commonly deciduous; petiole 3.5-10 cm long; blade chartaceous, ovate, unlobed, 7-27 x 3-20 cm, base truncate to deeply 

cordate. Pseudanthial inflorescences with moderately conspicuous green involucral bracts 10-15 x 4-6 mm; 7-10 

staminate flowers and 3 pistillate flowers. Staminate flowers 

with 8-22 stamens. Pistillate flowers with 6 ovate unlobed 

sepals; stylar column cylindrical, straight, 4-6 mm long, apex 

not dilated, stigma extending nearly full length of stylar 

column. Capsules much larger than any other species, 35-40 

mm in diameter. 

DISTRIBUTION.-Collected only near San Ignacio, Edo. 

Bolivar, Venezuela, near San Carlos de Rio Negro, Edo. 

Amazonas, Venezuela, and near Manaus, Brazil; five collections studied, from Venezuela and Brazil. 

ECOLOGY.-~CCUS in lowland wet forest below 500 m, at 

edge of forest or in forest canopy. The blue-green resin secreted 

by the inflorescence gland is collected by female Euglossa that 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

pollinate the flowers (Armbruster, 1988). Flowers and fruit 

present in December and January. 

SELECTED SPECIMENS EXAMINED.-Venezuela: Amazonas, 

15 km NE San Carlos de Rio Negro, Liesner 7471 (VEN). 

Brazil: Amazonas, Manaus, km 8, BR17, Rodrigues & Coelho 

1801 (INPA). 

NOTES.-This species has been found only in Guayanan and 

Amazonian Venezuela and Amazonian Brazil, but almost 

certainly occurs in Guyana. 

10. Dalechampia micrantha Poepp. 

Dalechampia micrantha Poepp., Nov. Gen. sp. pl., 3:19, 1841. [Type: Brazil, 

Megalosiylis poeppigii S. Moore, J. Bot., 54:208, 1916. [Type: Brazil, 

Amazonas, Ega (Tefe), Poeppig 2807 (holotype, W; isotypes, K, P).] 

Amazonas, Ega (Tefe), Poeppig (BM).] 

DESCRIPTION.-Twining vine reaching canopy. Leaves 

simple; stipules ovate, 2-3 x 1-2 mm, early deciduous prior to 

expansion of leaves; petioles 2-5 cm long; blades ovate, 

unlobed, 8-15 x 4-7 cm, base truncate, sometimes very 

shallowly cordate. Pseudanthial inflorescences with inconspicuous stipuliform involucral bracts 2-3 x 1.5-2 mm; 

usually 13 staminate flowers and 3 pistillate flowers. Staminate 

flowers with 4-9 stamens. Pistillate flowers with 5-6 unlobed 

sepals; stylar column clavate, slightly curved upwards, widest 

near blunt foveolate tip, stigma green, extending from tip to 

> 3/4 length of stylar column. Fruit a capsule, sepals deciduous 

prior to fruit maturation. 

DISTRIBUTION.-Venezuela (Bolivar), Surinam, Brazil 

(Amazonas), and Peru (Loreto); 20 collections studied, five are 

from Surinam. 

ECOLOGY.~CC~~S in moist to wet forests below 1000 m, 

forest edge or in secondary vegetation. Members of this species 

secrete a fragrance from the stigmatic surface of the pistillate 

flowers, attracting (in Venezuela) fragrance-collecting male 

Euglossa spp. (Hymenoptera: Apidae) that pollinate the 

flowers (Armbruster, unpublished observations). Flowers in 

late June (?), November through January, fruits present January 

to July. 

SPECIMENS EXAMINED.-surinam: SW of Lelydorp, Lindeman et al. 214 (U); Schomburgk 783 (P); Schomburgk 784 

(BM); Schomburgk s.n. (P); Jodensavanne-Mapane Creek, 

Wessels Boer 454 (K, NY, U). 

NOTES.-This species is a member of sect. Rhopalostylis and 

lacks the prominent resiniferous inflorescence gland characteristic of most of the genus. 

11. Dalechampia olympiana Kuhlman & Rodrigues 

Dalechampia olympiana Kuhlman and Rodrigues, Botanica, 5: 1, 1954. [Type: 

Brazil, Amazonas, Manaus, Rodrigues 225 (holotype, INPA).] 

DESCRIPTION.-Twining vine, stems pilose-lanate with 

orange hairs. Leaves simple; stipules lanceolate, 6-10 mm 

long, -2 mm wide; petioles 5-10 cm long; blades ovate, 

unlobed, 10-20 cm long, 6-13 cm wide, primary veins 

orange-hirsute, base ccordate, with narrow sinus. Pseudanthial 

inflorescences subsessile, with the two subequal, inconspicuous, green, stipuliform, hirsute, involucral bracts, - 12 staminate flowers, and 3 pistillate flowers. Staminate flowers with 

9-14 stamens. Pistillate flowers with 4-6 sepals, these hirsute, 

unlobed, lance-linear, 3.0-4.0 mm long, 0.5-1.5 mm wide; 

stylar column weakly clavate, straight, 6-10 mm long, basal 

portions hirsute, stigmatic region covering 1/2-3/4 of length. In 

fruit, pistillate pedicels 3-8 cm long, capsules hispid with 

yellow-brown hairs, -18 mm in diameter; seeds fattened 

spheroidal, 5.0-5.5 mm long, 4.3-4.5 mm high, surface 

smooth, mottled brown and gray or black and gray. 

DISTRIBUTION.-Apparently endemic to Amazonas and the 

Guianas; collected only in the Manaus region and recently in 

Guyana; five collections studied, of which one is from the 

Guianas (G: 1). 

EcoLoGY.-Pollination ecology is unknown. Flowers present in October; fruits present in May. 

SPECIMENS EXAMINED.-Guyana: Maburo region, Pibiri 

compartment, plot 7, 05'1.95'N, 58'37.73'W, Ek et al. 1187 

(U). 

NOTES.-An unusual member of section Rhopalostylis in 

having several, or more usually, numerous, subsessile pseudanthia in the axils of the leaves, and in having dense orange 

pubescence on the stems, petioles, and major leaf veins. Found 

in the Guianas in 1996; heretofore known only from the 

Manaus region of Amazonas, Brazil. 

12. Dalechampia papillistigma Armbruster 

Dalechampia papillistigma Armbruster, Brittonia, 41 :49, 1989. [Type: Venezuela, Bolivar, Santa Elena de Uairen, 4'36", 6lo15'W, Annbruster et al. 

87-155 (holotype, MO!; isotypes, ALA!, MY!, NY!, VEN!).] 

DESCRIPTION.-Twining vine reaching lower canopy. 

Leaves palmately compound, 3-5 leaflets; stipules lanceolate 

3-7 x 1-2 mm; petioles 2-10 cm long; blades chartaceous, 

that of center leaflet oblancelate to obovate, 4-15 x 2-7 cm, 

base cuneate; blades of lateral leaflets similar but lanceolate to 

ovate with asymmetrical base. Pseudanthial inflorescences with 

moderately conspicuous, spathulate, green involucral bracts 

5-8 x 3-4 mm; -10 staminate flowers and 3 pistillate flowers. 

Staminate flowers usually with 70-80 stamens. Pistillate 

flowers with 6 sepals, each with 2-4 longitudinal lobes; stylar 

column cylindrical, straight, 2.7-3.3 mm long, with distinct 

capitate tip; stigmatic surface densely papillate, confined to the 

swollen tip of style. 

DIsTRIBuTIoN.-Known only from southeastern Edo. Bolivar, Venezuela, near Brazilian border; likely occurs in northern 

Brazil and Southwestern Guyana. Endemic to region; two 

collections studied, both are from Venezuela. 

ECOLOGY.-In moist forest 800-1000 m, at edge of riparian 

forest in savanna, reaching at least lower canopy. The 

inflorescence gland of this species secretes resin as a pollinator 

reward. Flowers and young fruit present in December. 
NUMBER 86 25 

SPECIMENS EXAMINED.-venezuela: Santa Elena de Uairen, 

NOTES.-Not closely related to any other species that we 

4"37'N, 6lo7'W, Armbruster et al. 87-154 (ALA). 

have examined. 

13. Dalechampia parvibracteata Lanj. 

FIGURE 7A,B 

Dalechampia parvibracteata Lanj., Recueil Trav. Bot. Neerl., 31:463, 1934. 

[Type: Guyana, Upper Demerara River, Jenman 4088 (holotype, K isotype, 

US!).] 

DESCRIPTION.-Twining vine. Leaves simple, unlobed; 

stipules lanceolate, 3-7 x 1-2 mm; petioles 3-10 cm long; 

blades ovate, widest at about midpoint, 6-15 x 5-13 cm, base 

shallowly to deeply cordate, with U-shaped sinus, usually 

about as wide as deep. Pseudanthial inflorescences with 

inconspicuous, green, stipuliform, involucral bracts, 3-5 x 2-3 

mm, and 3 pistillate flowers. Pistillate flowers with 6 unlobed 

sepals, these ovate-linear to obovate-linear, -5 x 1-1.5 mm; 

stylar column clavate, straight, 5-9 mm long, with thin, 

elongated, pubescent stipe and expanded stigmatic region 

restricted to final l/3 to l/4 of length, tip rounded. Seeds 4.3-5.1 

mm in diameter, flattened spheroidal. 

DISTRIBUTION.-Known from Guyana, Venezuela (Bolivar), 

Brazil (Amazonas), and possibly Surinam; 16 collections 

studied, two are from Guyana. 

ECOLOGY.-Flowers present November to June. Fruits 

present middle April to August (middle April in Guyana). 

SPECIMENS EXAMINED.4uyana: Berbice-Corentyne Region, Baba-Grant Sawmill, -0.5 mi [0.8 km] above Cow Falls, 

McDowell and Gopaul2264 (ALA, US). 

NOTES.-A member of sect. Rhopalostylis, this species 

appears to be most closely related to D. heterobractea, from 

which it differs in having monomorphic involucral bracts and 

the stigma confined to the terminal l/4 of the style. Nearly all 

specimens from the Guianas assigned to this species actually 

belong to D. heterobractea. 

14. Dalechampia scandens L. 

Dalechampia scandens L., Sp. pl., 1053, 1753. [Type: West Indies, illustrated 

by Plumier, Plantarum Americanum, fasciculus 5, pl. 101 (original plate 

at P).] 

14a. Dalechampia scandens var. scandens 

FIGURE 8 

Dalechampia mollis Vahl, Eclog. amer., 3:44, 1807. [Type: Colombia, Santa 

Dalechampia ruboides Kunth in H.B.K., Nov. gen. sp., 2:102, 1817. [Type: 

Dalechampia sidaefolia Kunth in H.B.K., Nov. gen. sp., 2:102, 1817. [Type: 

Dalechampia guianensis Klotzsch, London J. Bot., 2:43, 1843. [Type: British 

Dalechampia brevipes Briq., Annuaire Conserv. Jard. Bot. Geneve, 4:609, 

Marta, von Rohr (C).]. 

Venezuela, Angostura, Humbolt & Bonpland (P).] 

Venezuela, Pararuma Island and Angostura, Humbolt & Bonpland (P).] 

Guiana, Schomburgk s.n.1 

1900. [Type: Colombia, Guanaguana, Funk 64 (G).] 

DESCRIPTION.-Twining or sprawling vine. Stems with 

sparse to dense, spreading, straw-colored to orangish pubescence. Leaves simple; stipules lanceolate to ovate, 3-8 x -2 

mm; petiole 2-8 cm long; blade chartaceous, ovate, usually 

3-lobed, rarely unlobed, 3-10 x 3-13 cm, apex of each lobe 

acuminate, leaf base shallowly to deeply cordate, glabrescent 

above, sparsely to densely pubescent below. Inflorescence 

pseudanthial, on peduncle 2-6 cm long; the two bracts 

involucral, pale green to greenish white at anthesis, broadly 

ovate, 1.5-3.0 x 1.5-3.0 cm, with 5 major veins, apex divided 

into three lobes to -l/z length of bract; inflorescence contains 

10 staminate flowers and 3 pistillate flowers; inflorescence 

gland secreting clear to whitish resin. Staminate flowers with 

25-35 stamens. Pistillate flowers with calyx of 8-12 sepals, 

these pinnatifid; pistil with elongate stylar column, 4-8 mm 

long, tip only slightly and asymmetrically dilated, -0.5 mm in 

diameter. Fruit a capsule; seeds spherical, mottled brown and 

gray, 3.5-4 mm in diameter, surface smooth. 

DISTRIBUTION.-Widely distributed from Mexico to Argentina; widespread in the Guianas; 70 collections studied, 39 of 

which are from the Guianas (G: 1 1, S:7, FG:2 1). 

ECOLOGY .-Clambering vine of open areas, secondary 

scrub, and margins of dry to moist forest below 1000 m. This 

species is pollinated by resin-collecting female euglossine bees 

(Euglossa, Eulaema), megachilid bees (Hypanthidium), and 

stingless bees (Trigona) (Armbruster, 1985). Flowers yearround. 

SELECTED SPECIMENS EXAMINED.-GUyana: Atkinson 

Field, Demerara, Harrison 1769 (K, NY); road from Karanambo to Yupukari, 3"45'N, 59"20'W, Maas et al. 7222 (B, 

US). Surinam: Zuid River, 2 km above confluence with Lucie 

River, 3"20'N, 56"49'W-3"10'N, 56"29'W, Irwin et al. 55831 

(NY, U, US); near Voltzberg, Saramacca, 4"41'N, 56"11'W, 

Webster 24143 (NY, U). French Guiana: Camopi River, 

Belvedere Mt., Granville 6940 (B, CAY, P); St. Laurent, Piste 

de Paul Isnard, Skog & Feuillet 7427 (CAY, NY, P, U, US). 

VERNACULAR NAMES.-Surinam: brongwirie (Surinamese), 

Lanjouw h Lindeman 1114 (NY, U), from label; palaramauloeloe (Carib), Lanjouw & Lindeman 1807 (NY, U), from label. 

NOTES.-This variety belongs to a highly variable species 

with numerous described infraspecific taxa and extensive 

ecotypic differentiation (Pax and Hofhann, 19 19b; Armbruster, 1985). The entire complex is in need of revisionary 

treatment. 

14b. Dalechampia scandens var. fallax Muell. Arg. 

Dalechampia scandans var. fallax Muell. Arg. in Martius, FI. bras., 11(2):658, 

1874b. [Type: Brazil, Par& Santarem, Spruce 199 (holotype, G).] 

DESCRIPTION.-Twining or sprawling vine. Stems with 

dense, spreading, orange pubescence. Leaves simple; stipules 

lanceolate to ovate, 4-10 x 2-3 mm; petiole 3-10 cm long; 

blade chartaceous, ovate, usually 3-lobed, rarely unlobed, 6- 15 

x 6-20 cm, apex of each lobe acuminate, leaf base deeply 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 8.-Dalechampia scandens L. var. scandens: A, habit; B, inflorescence; C, staminate flower; D, capsule 

(based on Pittier 10351). 
NUMBER 86 27 

cordate, sparsely pubescent above, densely tomentose below. 

Inflorescence pseudanthial, on peduncle 8-12 cm long; the two 

bracts involucral, white at anthesis, broadly ovate, 2.0-3.5 x 

2.0-3.5 cm, with 5 major veins, apex divided into three lobes 

to near middle of bract; margins with glandular tentacular 

processes 1 .O- 1.5 mm long; inflorescence gland secreting clear 

to whitish resin. Staminate flowers with 25-35 stamens. 

Pistillate flowers with calyx of 8-12 sepals, these pinnatifid; 

pistil with elongate, umbraculiform stylar column, 8-12 mm 

long, stigmatic surface confined to distal ‘/3 of column; tip 

distinctly expanded, symmetrically peltate, -2 mm in diameter. 

DISTRIBUTION.~CCWS in southern Guyana and perhaps 

eastern Brazil (see “Notes” below); two collections studied, 

both are from Guyana. 

SPECIMENS EXAMINED.4Uyana: Upper Takutu-Upper 

Essequibo Region, 5 km NNE of Dadanawa, Gillespie 19.55 

(ALA); by Rupununi River, Turukwan, Rupununi, Cook 44 

NOTES.-This taxon differs from D. scandens var. scandens 

in several characters, including the shape of the stylar column, 

capitate tentacular processes on the bracts, and the capitate 

glandular margins of the pistillate involucellar bract. It is 

tentatively assigned to D. scandens var. fallax. However, this 

material differs from D. scandens var. fallax (as described by 

Mueller (1874b) and Pax and Hoffmann (1919b)) in having 

larger involucral bracts and pistillate involucellar bracts with 

capitate glandular margins. The taxon treated here should 

probably be recognized as a distinct species, although it may 

not be conspecific with the type material of D. scandens var. 

fallax. Formal recognition of this taxon must await a 

revisionary study of the D. scandens species complex. 

w9 U). 

15. Dalechampia stipulacea Muell. Arg. 

Dalechampia stipulacea Muell. Arg., Linnaea, 34:221, 1865. [Type: Peru, 

Pavon (holotype, G).] 

DESCRIPTION.-Twining vine sometimes reaching lower 

canopy. Leaves simple; petiole 2-6 cm long; blade chartaceous, usually ovate, rarely lanceolate, usually 3-lobed, 5-12 x 

6-14 cm, apex of each lobe narrowly acuminate, leaf base 

deeply cordate, sparsely pubescent above and below, margins 

usually capitate glandular; stipules lanceolate-ovate, 10-20 x 

3-5 mm, margins capitate glandular. Inflorescence pseudanthial; the two bracts involucral, pale green at anthesis, broadly 

ovate, tip deeply 3-lobed, 4-6 x 2-4 cm, sparsely pubescent, 

margins usually capitate glandular; resiniferous gland secreting 

yellow resin. Staminate flowers with 30- -50 stamens. 

Pistillate flowers with calyx of 10-12 sepals, these pinnatifid, 

capitate glandular, hispid; pistil with elongate stylar column, 

10-15 mm long, tip moderately dilated. Fruit a capsule, surface 

glabrescent; seeds spherical, mottled brown and gray, -4 mm in 

diameter, surface smooth. 

DISTRIBUTION.-Widespread from western Venzuela, Colombia, and Peru, south to southern Brazil, Argentina, and 

Paraguay. Absent or rare in most of Amazonia and the Guianas. 

Locally distributed in French Guiana, thus far known only from 

Saul area; 15 collections studied, of which two are from French 

Guiana. 

ECOLOGY .-In open areas, including secondary vegetation 

and edges of moist to wet forest, below 1000 m elevation. The 

species is distinctive in secreting resin from vegetative parts; 

this resin inhibits feeding by both generalist and specialist 

insect herbivores and presumably plays a role in defense 

(Armbruster, unpublished observations). The inflorescence 

gland secretes copious yellow resin that is collected by 

pollinating female euglossine bees (Eulaema) in Venezuela. In 

French Guiana we saw only visitation by Hypanthidium sp. 

(Megachilidae), which are too small to be efficient pollinators. 

Flowers have been collected in August and June in French 

Guiana; it appears to flower year-round in Venezuela. 

SPECIMENS EXAMINED.-French Guiana: Route de Belizon 

between Eaux Claires and entrance to Boeuf-Mort trail, Mori et 

al. 23721 (CAY); -4 km north of Saul, Route de Belizon at 

junction with Boeuf Mort trail, Armbruster & Edwards 95-03 

(CAY, ALA). 

NOTES.-This species is distinctive in the Guianan region in 

having stipules, involucral bracts, and leaves with strongly 

capitate glandular margins. In some plants the leaf margins are 

eglandular, but even in these plants the stipules have glandular 

appendages. 

Dalechampia stipulacea has been only recently collected 

from the Guianas, from just one small area near Saul, French 

Guiana. Judging from collections, the species is generally 

absent from the Guianas and Amazonia. The nearest populations we are aware of are - 1500 km away in Brazil and - 1800 

km away in western Venezuela. Surprisingly, the Guianan 

material closely resembles the Venezuelan material. It is not 

known whether this apparent tremendous disjunction represents incomplete collecting in intervening regions, strange 

biogeographic history, or recent introduction (by miners?) in 

the Saul area. 

16. Dalechampia tilivolia Lam. 

Dalechampia tiliifolia Lam., Encycl., 2:257, 1786. [Type: possibly Peru, Herb. 

Jussieu (holotype, P).] 

Dalechampia peruviana Lam., Encycl., 2:257, 1786. [Type: Peru, Herb. 

Jussieu (P).] 

Dalechampia heterophylla Vahl, Eclog. amer., 3:44, 1807. [Type: Cayenne, 

Von Rohr (C).] 

Dalechampia pruriens Griseb., F1. Brit. W. I., 51, 1859. [Type: Trinidad, 

Crueger (GOET).] 

DESCRIPTION.-Twining vine sometimes reaching lower 

canopy. Leaves simple; stipules lance-linear, 3-7 x -1 mm; 

petiole 2-8 cm long; blade chartaceous, usually ovate, rarely 

lanceolate, usually 3-lobed mixed with unlobed when in bloom, 

6-15 x 4-13 cm, apex of each lobe acuminate, leaf base 

shallowly to deeply cordate, sparsely pubescent above, velutinous below. Inflorescence pseudanthial; the two bracts involu- 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cral, cream-white at anthesis, broadly ovate, tip shallowly 

3-lobed, 3-6.5 x 3-6 cm, velutinous, with 7-9 major veins; 

resiniferous gland secreting yellow resin. Staminate flowers 

with -40 stamens. Pistillate flowers with calyx of 10-13 

sepals, these pinnatifid, densely hispid; pistil with elongate 

stylar column, 6-12 mm long, tip asymmetrically dilated to 

umbraculiform. Fruit a capsule, surface sparsely to densely 

hispid; seeds spherical, mottled brown and gray, -5 mm in 

diameter, surface smooth. 

DISTRIBUTION.-Widespread from Mexico to southern Brazil. Widespread in unforested parts of the Guianas; 50 

collections studied, of which 33 are from the Guianas (G:l, 

FG:32). 

ECOLOGY.-In open areas, secondary scrub, and edge of dry 

to moist forest below 1800 m. The inflorescence glands secrete 

yellow resin that is collected by pollinating female euglossine 

bees (Eulaema, Euglossa) (Armbruster and Herzig, 1984). 

Flowers in the dry season in Mesoamerica; flowers collected in 

the Guianas throughout the year; hits mostly March to June. 

SELECTED SPECIMENS EXAMINED.4uyana: Iramaipang, 

Kanuku Mts., FD 5978 (K, NY). French Guiana: near Cayenne, 

Cremers 9429 (B, CAY, MO, NY, P, US); route to Stoupan, Ile 

de Cayenne, Feuillet 1754 (CAY, P); Mahury Mt., Rorota, 

Granville 260 (CAY, P, U); Mt. St. Martin, Cayenne Island, 

4"56'N, 52"17'W, Hoff5357 (B, CAY, NY, P, US); route to 

Stoupan, Ile de Cayenne, Oldeman B-802 (CAY, U); north of 

Saul, Prevost 1808 (CAY, U, US). 

NOTES.-Very closely related to D. affinis, from which it 

differs by having at least some 3-lobed leaves when in bloom, 

involucral bracts that are always shallowly 3-lobed and often 

nearly as wide as long, and style tips asymmetrically dilated 

rather than umbraculiform. 

17. Dalechampia aff. cissifolia Poepp. 

Dalechampia aff. cissifolia Poepp., Nov. gen. sp. pl., 3:20, 1841. [Type: Peru, 

Maynas, Yurimaguas, Poeppig 2085 (holotype, W; isotype, G).] 

DESCRIPTION.-Twining or sprawling vine. Leaves usually 

compound, rarely simple; stipules lanceolate, 3-5 x 0.5-1 .O 

mm; petiole 0.5-2.0(4.5) cm long; blade chartaceous, leaflets 

lanceolate, 2.5-6.5(8) x 0.5-2 cm, apex of each leaflet acute, 

central base cuneate, laterals oblique, glabrescent to sparsely 

pubescent above and below. Inflorescence pseudanthial, on 

peduncle 0.5-2.0 cm long; the two bracts involucral, leaf-green 

at anthesis, broadly ovate, 1.0-1.5 x 0.8-1.5 cm, with 5 major 

veins, apex entire or divided into three lobes to -'I4 to '12 length 

of bract; inflorescence contains -10 staminate flowers and 3 

pistillate flowers; inflorescence gland secreting resin. Pistillate 

flowers with calyx of 8-12 sepals, these pinnatifid; pistil with 

elongate stylar column, 1.5-3 mm long, tip only slightly and 

asymmetrically dilated. Fruit a capsule; seeds spherical, 

mottled brown and gray, -4 mm in diameter, surface smooth. 

DISTRIBUTION.-Material referable to this taxon has been 

collected only in Guyana; two collections studied, both are 

from Guyana. 

SPECIMENS ExAMINED.4uyana: Upper Takutu, Upper 

Essequibo Region, SE Kanuku Mts., Gillespie 1832; Upper 

Takutu, Upper Essequibo Region, near Surama village, 

McDowelI 203 7. 

NOTES.-The Dalechampia cissifolialtriphylla complex is 

badly in need of revision. Until such a revision is completed it 

is difficult to determine the taxonomic status of this taxon. It is 

almost certainly distinct at the species level from D. cissifolia. 

18. Dalechampia aff. triphylla Lam. 

Dalechampia aff. triphylla Lam., Encyl., 2:258, 1786. [Type: Brazil, Dombey 

(holotype, P [Herb. Jussieu 162921; isotype, NY).] 

DEsCRIPT1oN.-Twining or sprawling vine. Leaves usually 

compound, rarely simple; stipules' lanceolate, 4-7 x 1 .O-1.5 

mm; petiole 4-10 cm long; blade membranous to chartaceous, 

leaflets lanceolate, 4-10 x 1.5-3 cm, apex of each leaflet acute, 

central base cuneate, laterals oblique, glabrescent to sparsely 

pubescent above and below. Inflorescence pseudanthial, on 

peduncle 1-2 cm long; the two bracts involucral, leaf-green at 

anthesis, broadly ovate, 1.2-1.5 x 1.5-2.0 cm, with 5 major 

veins, apex divided into three lobes to -I13 to ]/2 length of bract; 

inflorescence contains -10 staminate flowers and 3 pistillate 

flowers; inflorescence gland secreting clear resin. Pistillate 

flowers with calyx of usually 6 sepals, these with broad rachis, 

width > *I4 length, and shallowly to deeply lobed, densely 

ciliated margins; pistil with elongate stylar column, 4-5 mm 

long, tip only slightly and asymmetrically dilated. Fruit a 

capsule; seeds spherical, mottled brown and gray, -3 mm in 

diameter, surface smooth. 

DISTRIBUTION.-Occurs in Surinam and French Guiana; 

four collections examined, all are from the Guianas (S:3, FG: 1). 

EcoLoGY.-Occurs in disturbed sites in lowland forests. 

Floral resin is collected in Surinam by female Hypanthidium cf. 

panamense (Megachilidae), which pollinate the flowers 

(Armbruster and Herzig, unpublished observations). Apparently flowers year-round. 

SPECIMENS EXAMINED.-French Guiana: St. Laurent, piste 

de Paul Isnard, Billiet & Jadin 1613 (CAY). Surinam: 

Lindeman et al. 655 (U); Brownsberg Nature Park, Webster & 

Mohr 24122 (U); Brownsberg Nature Park, Armbruster & 

Herzig 85-103 (ALA). 

NOTES.-The Dalechampia cissifolialtriphylla complex is 

badly in need of revision. Until such a revision is completed it 

is difficult to determine the status of this taxon, except that it is 

definitely not conspecific with D. triphylla. In many respects 

this taxon is intermediate between D. triphylla and D. 

cissifolia, but the nature of the pistillate sepals indicates closer 

affinity to D. triphylla. 

4. Genus Haematostemon (Muell. Arg.) Pax & K. Hoffm. 

Haematostemon (Muell. Arg.) Pax & K. Hoffm., Pflanzenr., IV.147.IX, 68:3 1, 

Astrococcus sect. Haematostemon Muell. Arg., Linnaea, 34: 158, 1865. [Type: 

1919a. 

Haematostemon coriaceus (Baill.) Pax & K. Hoffm.] 
W 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

DESCRIPTION.-Shrubs or small trees, monoecious, latex 

absent. Leaves simple, alternate, petiolate; stipules small, 

deciduous; blade pinnately veined, margin entire to serrate; 

glands absent. Inflorescence a racemose thyrse, axillary, 

bisexual; lower nodes with solitary pistillate flowers or 

bisexual cymes with central pistillate flower, upper nodes with 

unisexual cymes of staminate flowers, axis terminating in a 

single pistillate flower; bracts narrowly triangular, small, 

eglandular. Staminate flowers pedicellate; sepals 4, valvate; 

corolla and disc absent; stamens 4; filaments conical, dilated 

and connate at base, with dilated bases sometimes resembling 

an intrastaminal pseudodisc (e.g., H. coriaceous); pistillode 

absent. Pistillate flowers pedicellate; sepals 4 (or more on 

terminal flower); corolla and disc absent; ovary 3-locular, 

distinctly 3-lobed with carpels globose, verrucate; styles 

completely connate into massive, elongate cup-shaped structure with a narrow hollow center, apex truncate and very shortly 

3-lobed, with lobes notched and papillose. Fruit (known only in 

H. coriuceous) capsular?, distinctly 3-lobed, 3-carpellate. 

Seeds unknown. 

DISTRIBUTION.-TWO species, one in Guyana, one in 

Venezuelan Guayana. 

LITERATURE.-PaX and Hoffmann (1919a:31-32). 

1. Haematostemon guianensis Sandwith 

FIGURE 9 

Haemutostemon guianensis Sandwith, Kew Bull., 1950: 133, 1951. [Type: 

Guyana, Eagle Mountain, Fanshawe 1123 = FD 3859 (holotype, K!; 

isotypes, FDG!, K!, NY!, U!).] 

DESCRIPTION.-Tree to 6 m tall. Branchlets sparsely 

pubescent to glabrate, pubescent at tip. Petiole 3-7 mm long, 

pubescent; blade thinly subcoriaceous, oblanceolate, 1 1-26 x 

4-7.5 cm long, apex acuminate, base cuneate and very 

narrowly auriculate, glabrescent. Inflorescence 5- 15 cm long, 

slender, axes pubescent; pistillate flowers 2-5, solitary at basal 

nodes, staminate flowers numerous in condensed cymes above, 

single pistillate flower at apex of inflorescence axis; bracts 

1-1.5 mm long. Flowers pink-brown. Staminate pedicel 2-2.5 

mm long, sparsely pubescent; bud subglobose, 1-1.5 mm long; 

sepals ovate, 1.2-1.5 x -1 mm, apex acute, glabrate, inner 

surface vermcate-papillate; filaments 0.5-0.7 mm long, dilated 

to 0.2-0.3 mm wide at base, pointed at apex. Pistillate pedicel 

1-2.5 mm long, densely pubescent; sepals oblong-lanceolate, 

-1.5 mm long, outer surface sparsely pubescent; ovary 

distinctly 3-lobed, -0.8 x 1.5-2 mm, densely verrucate, 

glabrous; stylar column oblong-obovoid, 2.5-3.5 x -2 mm, 

truncate at apex with broad lobes bearing horn-like papillae at 

comers. Fruit unknown. 

DISTRIBUTION.-Endemic to Guyana, where it is known 

only from near Mahdia in central Guyana; three collections 

studied. 

ECOLOGY .-Forest understory tree collected in kakaralli 

(Eschweileru spp.)-clump wallaba (Dicymbe spp.) forest on 

escarpment slope on red lateritic soil. Flowers and young fruits 

reported in January and March. 

SPECIMENS EXAMINED.4Uyana: Mahdiana Mt., Potaro, 

Atkinson 1 I4 (BM); Mahdiana Mt., Potaro, Funshawe 2869 = 

FD 6016 (FDG, K, NY, U, US). 

NOTES.- Huemutostemon may be distinguished by its small 

tree or shrub habit, staminate flowers with four sepals and four 

stamens, and styles that are connate into a massive narrowly 

cup-shaped stylar column. Huemutostemon coriaceus (Baill.) 

Pax & K. Hoffm., the only other member of the genus, is 

endemic to the Rio Guainia area of Amazonas state, Venezuela. 

This species may be distinguished from H. guianensis by its 

smaller, narrowly ovate leaf blades and shorter inflorescences. 

The related monotypic genus Astrococcus, endemic to the 

upper Rio Negro region of Venezuela and Brazil, may be 

distinguished by its 4-segmented extrastaminal disc, ovoidurceolate stylar column, and conspicously homed carpels. 

5. Genus Plukenetia L. 

Plukenetia L., Sp. pl., 1192, 1753. [Type: Plukenetia volubilis L.] 

Apodandru Pax & K. Hoffm., Pflanzenr., IV.147.IX, 68:20,1919a. [Lectotype: 

Apodundru loretensis (Ule) Pax & K. Hoffm.] 

Elaeophora Ducke, Arch. Jard. Bot. Rio de Janeiro, 4:112, 1925. [Type: 

Elaeophora abutifoliu Ducke.] 

DESCRIPTION.-Lianas or twining vines, monoecious or 

rarely dioecious; latex absent. Leaves simple, alternate, 

petiolate; stipules small, deciduous; blade pinnately or palmately veined, margin subentire to sermlate; l to several pairs 

of flat laminar glands present near base on adaxial blade 

surface; scattered laminar glands often present on abaxial blade 

surface, rarely present on adaxial surface (only in P supraglandulosa). Inflorescence a racemose thyrse, axillary or terminal 

on short shoot, bisexual with pistillate flower(s) at base and 

staminate flowers above in condensed cymes or less often 

unisexual; bracts triangular, small, eglandular. Staminate 

flower pedicellate; sepals 4-5, valvate; corolla absent; disc 

interstaminal, segmented or annular, or often absent; stamens 

15-40, free; filaments short to elongate or anthers sessile; 

pistillode absent. Pistillate flower pedicellate; sepals 4; corolla 

absent; disc absent; ovary 4-locular, 4-angled to deeply 

4-lobed, carpel lobes carinate or with tubercle or horn; styles 

partly to completely connate, stylar column cylindrical to 

globose or obovoid. Fruit a 4-seeded capsule or berry, 

subglobose to deeply 4-lobed; carpels carinate or with central 

tubercle or horn. Seeds subglobose, ovoid, or lenticular and 

often laterally compressed, ecarunculate, surface smooth or 

roughened. 

DISTRIBUTION.-Seventeen species, 12 in the Neotropics, 

four in Africa and Madagascar, one in Asia; five species in the 

Guianas. 

LITERATURE.-Pax and Hoffmann (1919a: 1 1-24); Gillespie 

(1993b). 
NUMBER 86 31 

Key to the Species of Plukenetia 

1. Leaf blades pinnately veined, elliptic in shape, acute, obtuse, or rounded at base; 

scattered laminar glands present on lower surface of blade ........... 2 

Leaf blades palmately veined or 3-veined at base, ovate or cordiform in shape, 

cordate, truncate, or rounded at base; laminar glands absent on lower surface of 

blade (or very small and marginal in P. verrucosa) ............... 3 

Stylar column slender-cylindrical, 2-4 mm long; ovary and capsule with pointed 

horn on each carpel; androecium consisting of 15-25 anthers sessile on globose 

receptacle; scattered laminar glands absent on upper surface of leaf blade .... 

................................... 1. €? loretensis 

Stylar column stout-cylindrical, -1.6 mm long; ovary and capsule with rounded 

tubercle on each carpel; androecium consisting of outer whorl of 4 stamens with 

filaments, and inner cluster of 6-9 anthers sessile on globose receptacle; 

numerous scattered laminar glands present on upper surface of leaf blade .... ............................... 3. €? supraglandulosa 

Styles completely connate into subglobose or obovoid column; stamens in outer 

whorl with filaments, inner anthers sessile; fruit 4-lobed, less than 1.5 cm in 

diameter, dehiscent ........................ 4. €? verrucosa 

Styles connate into cylindrical column, free at apex; all stamens with filaments; fruit 

Stylar column 15-30 mm long, arms -1 mm long; fruit 4-lobed, 2.5-4(-6) cm in 

diameter, dehiscent; staminate bud subglobose; filaments conical, -0.5 mm long; 

leaf base cordate or truncate .................... 5. I? volubilis 

Stylar column 3-7 mm long, arms 1.5-2.5 mm long; fruit subglobose, squarish in 

cross-section, 6-1 1 cm in diameter, indehiscent; staminate bud narrowly 

oblong-ellipsoid; filaments slender, 2-3 mm long; leaf base rounded or obtuse 

2. 

3. 

greater than 2.5 cm in diameter .......................... 4 

4. 

................... 

1. Plukenetia loretensis Ule 

Plukenetia loretensis Ule, Verh. Bot. Vereins Prov. Brandenburg, 81, 1908. 

Apodandra loretensis (Ule) Pax & K. Hoffm., Pflanzenr., IV.147.IX, 68:21, 

[Type: Peru. Loreto. Iquitos, Ule 6837 (isotype, G!).] 

1919a. 

DESCRIPTION.-Vine or liana, monoecious. Young stems 

tomentose, becoming sparsely pubescent, often twining. Petiole 0.5-2 cm long, tomentose; blade chartaceous, elliptic, 

6-12(-16) x 3-7(-9) cm, apex acuminate, base acute to 

obtuse and shortly decurrent, margin sermlate, glabrescent to 

sparsely pubescent on major veins above and below, pinnately 

veined; basilaminar glands 2-4 pairs, circular, -1 mm in 

diameter, or sometimes confluent into a single elongate pair, -3 

mm long, on decurrent leaf base; scattered laminar glands 

circular, -0.5 mm wide, near margin. Inflorescences axillary, 

racemose, 1-6 cm long, axes sparsely to densely tomentose; 

pistillate flower single at base, staminate flowers in condensed 

cymes above; bracts ovate-triangular, -1 mm long, sparsely 

pubescent. Staminate pedicel 4-8 mm long, tomentose; bud 

subglobose, -1.5 mm long, apex obtuse; sepals 4, ovateelliptic, 2-2.5 mm long, sparsely pubescent; anthers 15-25, 

sessile on globose receptacle. Pistillate pedicel 2-3 mm long, 

tomentose; sepals triangular, -0.7 mm long, sparsely pubes- 

.. ............. .2. l? polyadenia 

cent, appressed to ovary; ovary 4-horned, 1-1.5 x 2-3 mm, 

densely tomentose except for horns; style cylindrical, 2-4 x 

-0.5 mm, sparsely tomentose; stigma 4-lobed, -1 mm wide. 

Capsule 4-lobed, 0.5-0.7 x 1-1.2 cm, puberulent, dehiscent, 

each carpel lobe with pointed horn, 3-6 mm long. Seeds 

subglobose to broadly lenticular, 5-6 x 3.5-5 x 4.5-5.5 mm, 

mottled pale and dark brown with course dark brown raised 

reticulations. 

DIsTRIBUTIoN.~olombia (Amazonas), Venezuela (hazonas, Bolivar), Guyana, Peru (Loreto, San Martin), Bolivia 

(Beni), and Brazil (Amazonas, Mato Grosso, RondBnia); in 

Guyana known only from the New River area; 53 collections 

studied, of which one is from Guyana. 

ECOLOGY.---Occurs in moist to wet lowland forest, below 

700 m. The single flowering and fruiting collection known 

from Guyana was made in September. 

SPECIMENS ExAMINED.-Guyana: Kanasenay, Guppy 203 = 

FD 71 79 (NY-3 sheets). 

VERNACULAR NAME.4uyana: Chichinat (FD 71 79). 

2. Plukenetia polyadenia Muell. Arg. 

FIGURE 10~-D 

Plukenetiapolyadenia Muell. Arg. in Mart., F1. bras., 11(2):334, 1874a. [Type: 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 10.-Plukenetia polyadenia Muell. Arg. and P volubilis L. A-D, P polyadenia: A, habit, showing 

axillary staminate inflorescences; B, pistillate inflorescences in leaf axils; C, staminate flower; D, pistillate flower 

(A,C based on Fanshawe 3383 = FD 6947 NY; B,D based on Black 52-141 73, P); E, P volubilis: E, pistillate 

flower (based on Zarucchi 2378, US). 
NUMBER 86 33 

Brazil, Prov. Maynas, Poeppig 2385 (holotype, W, photo F32544!, fragment 

at F!; isotype, G!).] 

Elaeophora polyadenia (Muell. Arg.) Ducke, Arch. Jard. Bot. Rio de Janeiro, 

5:146, 1930. 

Elaeophora abutifolia Ducke, Arch. Bot. Rio de Janeiro, 4:112, 1925. 

[Syntypes: Brazil. near Belem, Ducke HJBR 17892 (not seen); Tajapani R., 

Ducke HJBR 17893 (isosyntype, P!); Xingu R., Kuhlmann HJBR 17895 

(isosyntype, U!).] 

Plukenetia abutifolia (Ducke) Pax & K. Hoffm., Pflanzenfam., ed. 2, 19c:141, 

1931. 

DESCRIPTION.-Liana climbing to canopy, dioecious or 

sometimes monoecious. Branchlets glabrous to sparsely puberulous, often twining. Petiole 1.5-5 cm long, glabrous; blade 

chartaceous to subcoriaceous, elliptic to ovate-elliptic, 7- 1 1 

(-14) x 4-7(-10) cm, apex obtuse-acuminate with acumen 

slender 0.6-1.5 cm long, base rounded to obtuse, glabrous, 

3-veined at base; basilaminar glands 1 pair (rarely 2 pairs), 

transversely elliptic, -1.5-2 mm wide, with small knob 

between; scattered laminar glands absent. Inflorescences 

axillary, racemose or sometimes paniculate, usually unisexual, 

axes puberulous; staminate inflorescence 5-25 (rarely to 35) 

cm long, flowers numerous on scorpioid or dichasial-scorpioid 

cymose lateral branches 0.2-2 cm long, pistillate flowers 

occasionally present at basal-most 1-2 nodes on an otherwise 

staminate inflorescence; pistillate inflorescence 1.5-3 (rarely 

to 7) cm long, flowers 3-4(-lo), 1 or rarely 2 per lateral cyme 

branch, these 1-3 mm long; bracts triangular, -1 mm long, 

apex acuminate. Staminate pedicel 2-4 mm long; bud narrowly 

oblong-ellipsoid or oblong-ellipsoid, -3 mm long, apex acute; 

sepals 4, narrowly oblong, 3-4 mm long, apex acute, glabrous, 

reflexed at anthesis; stamens 20-25 on convex scaly receptacle; filaments 2-3 mm long, slender, attenuate at apex. 

Pistillate pedicel 0.5-1 mm long, glabrous to sparsely 

puberulent; sepals 4, triangular, 0.5-1 mm long, glabrous, 

appressed to ovary; ovary 4-angled, -1.5 x 2 mm, not distinctly 

winged or lobed, glabrous; styles connate into a narrow-conical 

column, 3-7 mm long, free at apex, arms 1.5-2.5 mm long, 

divergent. Infructescence up to 8 cm long; berry subglobose, 

quadrangular in cross section, each angle distinctly keeled, 

6-10 cm long x 6-1 1 cm in diameter, glabrous, fleshy, 

indehiscent, with broad conical apical protuberance -0.5 cm 

long. Seeds ovoid, 4.9-5.6 x 3.3-3.7 x 3.0-3.5 cm, apex 

narrowed, base rounded, dull brown, surface rough. 

DISTRIBUTION.-Widespread in the Guianas, eastern Venezuela, and the Amazon basin in Ecuador, Peru, Bolivia, and 

Brazil; 46 collections studied, of which 15 are from the Guianas 

(G:6, S:3, FG:6). 

EcoLoGY.4anopy liana in moist to wet lowland or lower 

montane forest, sea level to -1000 m. Flowers reported 

between February and August in the Guianas, fruits in August 

and October. Seed germination is epigeal and cryptocotylar, 

i.e., the cotyledons of seedlings are above ground but remain 

hidden within the seed coat (Gillespie, unpublished observations). 

SPECIMENS EXAMINED.auyana: Demerara, introduced 

from Fort Island, Beckett & Kortright 8640 (BRG, K, U); 

Camaria Falls, Cuyuni R., Fanshawe 3383 = FD 6947 (FDG, 

K, NY); Mt. Ayanganna, Tiller et al. 44950 (MO, NY, US); Mt. 

Ayanganna, Tillet et al. 44954 (NY); Kamarang R., Tillet & 

Tiller 45834 (NY, US); Oko Creek, Cuyuni R., Tutin 359 (K, U, 

US). Surinam: Emmaketen, Daniels & Jonker 1109 (US); 

along road near Moengo tapoe, Lindeman 6134 (AAU, DAV, 

F, K, U); Jodensavanne-Mapane Kreek area, Schulz 7284 (MO, 

U, US). French Guiana: vieille route de St. Laurent-Saut 

Sabbut, 10 Apr 1971, Fouque s.n. (CAY); Saul, Granville 

B.4413 (CAY); Saul, Grenand 647 (CAY); Saul, Moretti 461 

(CAY); Saul, Mori 15673 (NY, fruit only); Saul, Mori & 

Gracie 18668 (CAY, K, US). 

USES.-According to Ducke (1925) seeds are sold for their 

oil in local markets in the Brasilian state of Para. 

NOTES.-Ducke (1925, 1930) considered the species to be 

dioecious with dimorphic unisexual inflorescences in contrast 

to the bisexual inflorescences of all other species in the genus. 

The majority of collections examined are either staminate or in 

fruit. Two collections have pistillate flowers only, whereas 

three collections have bisexual inflorescences (e.g., Schultz 

7284) (Gillespie, 1993b). Therefore, the species appears to be 

at least sometimes monoecious. The above description of 

pistillate inflorescences was taken from two Brazilian collec - 

tions (Black 52-14137; Ducke 17893). 

3. Plukenetia supraglandulosa L. J. Gillespie 

FIGURE 11 

Plukenetia supraglandulosa L.J. Gillespie, Syst. Bot., 18:589, figs. 2, 1, 8, 10, 

1993. [Type: French Guiana, Sommet Tabulaire, zone centrale, versant 

occidental, about 40 km SE de Saul, Granville 3626 (holotype, US!; 

isotypes, CAY!, U!).] 

DESCRIPTION.-Liana, monoecious. Branches slender, branchlets puberulous. Petiole 0.5-2.0 cm long, puberulous; blade 

chartaceous, elliptic or ovate-elliptic, 7-13 x 3-8 cm, margin 

sermlate, apex slender-acuminate with acumen 0.5- 1 cm long, 

base acute, glabrescent except lower midrib puberulous, major 

veins and midrib sometimes sparsely puberulous below, 

pinnately veined, secondary veins distinctly looped; basilaminar glands in 1-3 pairs, circular or elliptic; laminar glands 

circular, on both surfaces, 10-20 near margin on abaxial 

surface, 8- 18 scattered between midrib and margin on adaxial 

surface. Inflorescences axillary, racemose, 2-8 cm long, axes 

puberulous, green; pistillate flower single at base, staminate 

flowers numerous in very condensed cymes above; bracts 

triangular, -1 mm long. Staminate pedicel 3.5-4.5 cm long, 

densely puberulous; bud subglobose, -1 mm long or less, apex 

rounded or broadly obtuse; sepals 4, elliptic, 1-1.3 mm long, 

apex acute and thickened, reflexed; disc 4-lobed, located 

between outer stamen whorl and inner stamen cluster; 

androecium of 4 outer stamens with filaments 0.3-0.4 mm long 

and 6-9 sessile anthers on globose receptacle. Pistillate pedicel 

6-12 mm long, puberulous; sepals 4, triangular, -1 mm long; 

ovary 4-winged, -1 x 2 mm, densely puberulous, wings 

rounded; style stout-cylindrical, -1.6 x 0.6 mm, sparsely 

puberulous. Capsule 4-lobed, -7 x 15 mm, sparsely puberulous, dehiscent, pendant, each carpel lobe with single blunt 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 86 35 

FIGURE 1 1 (left).-Plukenetia supraglandulosa L.J. Gillespie: A, habit; B, leaf 

blade base, adaxial surface, showing pair of basilaminar glands; C, leaf blade, 

abaxial surface, showing scattered laminar glands; D, young inflorescence in 

leaf axil, showing basal position of pistillate flower; E, upper part of 

inflorescence showing staminate flower buds in condensed cymes; F, staminate 

flower bud beginning to open; G, staminate flower at anthesis; H, pistillate 

flower; I, immature capsule; J, seed, showing dorsal, ventral, and lateral views, 

respectively (A-C,F,G,I based on Granville 3626, CAY, US; D,E,H based on 

Cowan 38204, US; J based on Granville et al. 10783, CAY). 

horn-like tubercle. Seeds lenticular, circular to broadly elliptic 

in outline, 7-7.3 x 4.6-4.8 x 6.8-7.1 mm, reddish brown with 

raised reticulate pattern. 

DISTRIBUTION.-Endemic to Surinam, French Guiana, and 

the Brasilian state of Amapa; five collections studied, of which 

one is from Surinam, three are from French Guiana, and one is 

from Amapa, Brazil. 

ECOLOGY .-Occurs in submontane forest on lateritic plateau 

summits at 500-800 m in French Guiana. Flowers reported in 

January, August, and November; fruits reported in January and 

August. 

SPECIMENS EXAMINED.-Surinam: Tumuc Humac Mts., 

Talouakem, 2’31‘N, 54’45’W, Acevedo 6022 (NY, US). French 

Guiana: Massif du Decou Decou, E summit, Granville 5315 

(U); Mont Atachi Bacca, S of summit plateau, 12 km SE of 

Gobaya Soula, 3”33’N, 53’55‘W, Granville et al. 10783 (CAY, 

US). 

4. Plukenetia verrucosa Smith 

FIGURE 12 

Plukenetia verrucosa Smith, Nova Acta Regiae SOC. Sci. Upsal., 6:4, 1799. 

[Type: Surinam, Herb. Linnaeus (holotype, LINN, microfiche IDC 

5073.1489.2!).] 

Plukenetia integrifolia Vahl, Eclog. amer., 3:43, 1807. [Type: Guyana, 

“Demerari,” von Rohr s.n. (holotype, C, photo, A!).] 

DESCRIPTION.-Twining vine, monoecious. Stems slender, 

glabrescent to sparsely pubescent; young stems sparsely to 

densely pubescent. Petiole 1-3 cm long, slender, pubescent; 

blade chartaceous, ovate or oblong-ovate, 5-12 x 2-7 cm, apex 

long-acuminate with acumen 0.5-1.5 cm long, base cordate, 

truncate-cordate or truncate with broad, usually shallow sinus, 

sparsely pubescent on margin and veins above and below, 

3-veined at base; basilaminar glands paired, subcircular to 

laterally elongate, 0.9-1.3 mm wide, with pair of stipels or 

small knob between; laminar glands circular, -0.3 mm wide, 

adjacent to margin. Inflorescence axillary, terminal on short 

shoot 1-4 cm long, or terminal and appearing leaf-opposed, 

racemose, 0.5-4 cm long, axes tomentose; pistillate flower 

single at base, staminate flowers numerous in condensed cymes 

above; bracts triangular, 0.5- 1 mm long, pubescent. Staminate 

pedicel 2-4.5 mm long, pubescent; bud globose, 1 - 1.4 mm in 

diameter; sepals 4, ovate, -1 mm long, apex acute; annular disc 

between outer stamen whorl and inner anthers; androecium of 

6-10 sessile anthers on a stout cylindrical receptacle and 4 

outer stamens with filaments -0.3 mm long. Pistillate pedicel 

2-4.4 mm long, tomentose; sepals 4, lanceolate, - 1.5 mm long, 

pubescent; ovary deeply 4-lobed, -1 x 2-2.5 mm, glabrous 

except for scattered hairs on lobes, lobes with prominent 

tubercle; style subglobose or obovoid, 1.5-2(-2.5) mm long 

and wide, glabrous, often with prominent 4-lobed stigmatic 

surface. Fruiting pedicel 0.8-1.5 cm long; capsule 4-lobed, -6 

x 11-14 mm, dehiscent, each carpel lobe prominently 

tuberculate. Seeds lenticular, subcircular in outline, -6 x 4 x 5 

mm, brownish gray with white markings, surface pitted. 

DISTRIBUTIoN.-Guianas, Trinidad, and northern Amazonian Brazil (Amapa, northeast Amazonas, Para, and Roraima); 14 collections studied, of which seven are from the 

Guianas (G: 1, S:3, FG:3). 

ECOLOGY.-Uncommon in disturbed areas or forest edge of 

lowland wet forest. Flowers reported in January, March, and 

July; fruits reported in January, July, and August. 

SPECIMENS ExAMINED.4Uyana: Camp Macaw Falls, 

Waini R., Beckett 8455 (BRG, NY). Surinam: Nickerie R., near 

Awawara, BW 893 (K, U); Jodensavanne, LBB 8565 (U); 

Voltzberg, Pulle 221 (U). French Guiana: St. Jean du Maroni, 

Benoist 893 (P); without specific locality, 1863, Melinosie (?) 

(P); Karouany (or Maroni), Sagot 22 (BM, G-BOIS, G-DC, K, 

MPU, P, U; consists of several collections). 

5. Plukenetia volubilis L. 

FIGURE IOE 

Plukenetia volubilis L., Sp. pl., 1192, 1753. [Type: West Indies. Illustration t. 

13 (lower ha14 in Plumier, Nov. pl. amer., 47, 1703.1 

Plukenetia peruviana Muell. Arg., Linnaea, 34: 157, 1865. [Syntypes: Peru, 

Herb. Pavon (G-DC!, G!); Peru, “prov. Maynas,” Poeppig 2110 (not seen).] 

Plukenetia macrostyla Ule, Verh. Bot. Vereins Prov. Brandenburg, 80, 1908. 

[Type: Brazil, Amazonas, Rio Jurua, bei Jabuni, Ule 5864 (isotype, G!, 

fragment at F!).] 

DESCRIPTION.-Twining vine or slender liana, monoecious. 

Stems glabrescent to pubescent; young stems pubescent. 

Petiole 2.5-7.5 cm long, glabrous to sparsely pubescent; blade 

chartaceous or membranous, triangular-ovate, 7- 13(-18) x 

4-lo(-18) cm, apex long-acuminate with acumen 1-2 cm 

long, base truncate to cordate, glabrescent below, 3-veined at 

base; basilaminar glands paired, circular to laterally elongate, 

1.5-3 mm wide, usually with small knob between; scattered 

laminar glands absent. Inflorescence axillary or terminal on 

short shoot, racemose, 5-18 cm long, axes sparsely to 

moderately pubescent; pistillate flower(s) 1 or rarely 2 at basal 

node(s), staminate flowers numerous in condensed cymes 

above; bracts narrowly triangular, 1.5-2.5 mm long. Staminate 

pedicel 2-3 mm long; bud subglobose, -2 mm long, apex 

rounded to obtuse; sepals 4, ovate-elliptic, -2 mm long, apex 

acute, glabrous; disc absent; stamens 20-25(-28) on convex 

receptacle; filaments -0.5 mm long, dilated at base, attenuate at 

apex. Pistillate pedicel -1.5 mm long; sepals 4, lineartriangular, 1.5-4 mm long, glabrous; ovary 4-winged, 1.5-2.5 

x 3-5 mm, glabrous; style column elongate-cylindrical, 15-30 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 86 37 

FIGURE 12 (left).-Plukenefia verrucosu Smith; A, habit; B, leaf blade base, 

adaxial surface, showing pair of basilaminar glands and pair of stipels; C, 

inflorescence, terminal but appearing leaf-opposed, with single basal pistillate 

flower; D, staminate flower showing two outer stamens having short filaments, 

an annular disc, and a central cluster of sessile anthers; E, pistillate flower; F, 

capsule, dried (A,B based on Sugot 22, P; C,D based on Prance et al. 11255, 

DAV; E based on Irwin 48796, US; F based on Burchell9369, K). 

x 0.7-1.5 mm, 4-lobed at apex, arms -1 mm (rarely to 3 mm) 

long, often curled adaxially. Capsule deeply 4-lobed, 1.5-2.5 x 

2.5-4(-6) cm, glabrous, initially fleshy, becoming woody and 

dehiscent, each carpel lobe with central wing to 2 mm wide. 

Seeds lenticular, broadly oblong in outline, 1.6-2 x 0.7-0.9 x 

1.5- 1.7 cm, brown with course dark brown markings. 

DISTRIBUTION.-Widespread in the Lesser Antilles and 

South America, where it is found primarily in the northern and 

western regions and margins of the Amazon Basin in Surinam, 

Venezuela (Amazonas), Colombia (Meta), Ecuador, Peru, 

Bolivia, and Brazil (western Amazonas, Para); 50 collections 

studied, of which two are from Surinam (S:2). 

EcoLoGY.4ccurs in disturbed areas or forest edge of 

lowland moist or wet forest, below 900 m. In Surinam flowers 

reported in April and August, fruits in August and September. 

SPECIMENS EXAMINED.--SUnnam: West River, 2-5 km 

SW of Juliana Top, Irwin et al. 54897 (F, GH, MO, NY, U, 

US); area of Kalalebo Dam project, Lindeman et al. 636 

UsEs.-vines are occasionally cultivated in western Amazonia for the seeds that apparently are edible when roasted 

(although seeds of wild plants were found to be distasteful and 

(MO, U). 

inedible when eaten raw). Cultivated plants frequently have 

larger 5-6 carpelled fruit and more prominently serrate leaf 

margins (Gillespie, 1993b). 

6. Genus Tragia L. 

Trugiu L., Sp. pi., 980, 1753. [Lectotype: I: volubilis L.] 

DESCRIPTION.-Herbs, twining vines or subshrubs, monoecious or rarely dioecious. Stems and foliage usually covered 

with urticating hairs. Leaves simple, alternate, petiolate; 

stipules usually small, deciduous; blade pinnately or palmately 

veined, entire, lobed, or serrate; glands absent. Inflorescence a 

racemose or little branched paniculate thyrse, axillary, terminal 

on short shoots, or terminal but appearing leaf opposed, 

bisexual or rarely unisexual, with pistillate flower(s) basal; 

bracts small, triangular, eglandular. Staminate flowers pedicellate; sepals 3-6; corolla absent; disc intrastaminal and 

subtriangular or extrastaminal and segmented into filiform 

segments, often reduced or absent; stamens (1 -)2-50; filaments present or anthers sessile; pistillode absent. Pistillate 

flowers pedicellate; sepals (3-)6; corolla absent; disc absent; 

ovary 3-locular, densely covered with urticating hairs; styles 

slender, partly connate. Fruit a 3-seeded capsule, 3-lobed, 

dehiscent, usually covered with urticating hairs. Seeds globose 

to ellipsoid, ecarunculate. 

DISTRIBUTION.-About 130 species in tropical and subtropical regions of the New World, Africa, south and southwestern 

Asia, and Australia; four species are in the Guianas. 

LITERATUFS-P~X and Hoffmann (1919a:32-101); Gillespie (1 994b). 

Key to the Species of Tragia 

1. Inflorescence appearing dichotomous, consisting of staminate main axis to 18 cm 

long and a pistillate basal branch to 10 cm long; stamens 8-17; staminate disc 

segments present; fruit pedicel 1-4 mm long .................. 2 

Inflorescence racemose, to 10 cm long, with 1-2 pistillate flower(s) at the basal 

node(s); stamens 1-3; staminate disc segments absent; fruit pedicel 1-40 mm 

long ........................................ 3 

Leaf blade palmately veined, base cordate; stamens 7-10; disc segments 6-7 

.................................... 1. T fendleri 

Leaf blade pinnately veined, base acute or obtuse; stamens 14-17; disc segments 

8-9 ................................ 2. T lessertiana 

Glandular hairs present on stem, petioles, and inflorescence axes; inflorescence 4-9 

cm long; fruit pedicel less than 0.5 cm long ........ 3. T tabulaemontuna 

Glandular hairs absent; inflorescence 1-4 cm long; fruit pedicel 1.5-4 cm long 

................................... .4. T volubilis 

2. 

3. 

1. Tragia fendleri Muell. Arg. 

FIGURE I3A-D,G 

Brazil, Alto Amazonas, Maribi, Japurensi District, Martius s.n. (holotype, 

M, photo F6626!).] 

DEsCRIPTION.-Twining vine, monoecious. Stems slender, 

covered with urticating hairs. Leaves covered with urticating Trugiu fendleri Muell. Arg., Linnaea, 34: 179, 1865. [Type: Venezuela, near 

~i~~~i~~, 1208 (holovpe, G-DC!; isotypes, G!, G-DC!, GH!, MO!).] .. .. 

Trugiu japurensis Muell. Arg. in Mart., FI. bras., 11(2):404, 1874b. [Type: hairs; petiole 2.5-1 1 cm long; blade chartaceous, elliptic, ovate 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE I3.-Tragia fendleri Muell. Arg. and Tragia lessertiana (Bail].) Muell. Arg. A-D,G, T fendleri: A, habit; 

B, inflorescence showing staminate main axis and pistillate branch at left; c, pistillate flower; D, staminate flower; 

G, capsule (A-D based on Gillespie et al. 1678, US; G based on Steyermark et al. 114589, MO); I: lessertiana: 

E, pistillate flower; F, staminate flower (E,F based on Webster 241 19, DAV). 
NUMBER 86 39 

or oblong-ovate, 8-15.5 x 3.5-8.5 cm, margin dentate with 

teeth large and somewhat rounded, sometimes lobate near base, 

apex long-acuminate, base cordate, sinus angular, narrow to 

moderately broad, 0.6-1.6 cm deep, palmately veined with 5 

basal veins. Inflorescence terminal, consisting of a staminate 

main axis with a single pistillate basal branch; peduncle 1-3 

cm long, elongating in fruit to 6.5 cm; staminate axis racemose, 

1.5-7 cm long; pistillate axis racemose, 2.5-7 cm long, 

elongating in fruit to 17 cm; bracts lanceolate, apex attenuate, 

staminate bracts -2 mm long, pistillate bracts 3-4 mm long; 

flowers numerous, 1 per node; axes densely covered with 

urticating hairs. Staminate pedicel 2.0-3.5 mm long; bud 

obovoid, 2.0-2.6 mm long; sepals 3(-4), oblong elliptic or 

elliptic, -2.5 mm long, apex acute; disc segments 6-7, slender, 

ligulate, 0.7-1 mm long, apex narrowed; stamens 7-10, outer 

whorl of 6(-7) alternating with and basally adnate to disc 

segments, inner cluster of 2-3; filaments slender, 1-1.5 mm 

long; anthers sagitate, -1 mm long. Pistillate pedicel 0.3-0.5 

mm long; sepals 6, lanceolate, 3-3.5 mm long; ovary globose, 

- 1 mm in diameter, very densely covered with urticating hairs; 

styles 3-3.5 mm long, connate one-third to one-half of length, 

inner surface papillate. Fruiting pedicel 1.5-4 mm long; sepals 

persistant, 5-8 mm long; capsule distinctly 3-lobed, 3-5 x 

6- 10 mm, densely covered with urticating hairs. Seeds globose 

or ellipsoid-globose, 3-3.5 mm long, pale brown with 

red-brown spots. 

DISTRIBUTION.-Panama, Venezuela, Guyana, and Brazil 

(Amazonas); nine collections studied, of which two are from 

Guyana. 

ECOLOGY.4ccurs in Guyana on white sand savanna or at 

the savanndseasonal forest margin. In Guyana flowers reported 

between April and June; fruits reported in June. 

SPECIMENS ExAMINED.--Guyana: Natun Bush Island, -2 

km SSW of Dadanawa, Rupununi Savannas, Gillespie et al. 

1678 (US); NW slopes of Kanuku Mts. in drainage of 

Moku-moku Creek, Smith 3558 (A, F, NY, U, US). 

2. Tragia lessem'ana (Baill.) Muell. Arg. 

FIGURE 13~,~ 

Tragia lesserriana (Baill.) Muell. Arg., Linnaea, 34: 178, 1865. 

Bia lessertiana Baill., Etude Euphorb., 502, 1858. [Type: Brazil, Herb. 

Delessert (holotype, G!; fragments at F!, P!).] 

DESCRIPTION.-Twining vine, monoecious. Stems slender, 

often woody, sparsely covered with urticating hairs. Petiole 

1-5 cm long, sparsely covered with urticating hairs; blade 

chartaceous, oblanceolate or narrowly elliptic, 6-1 1.5 x 2-5.5 

cm, margin serrate, apex acute or acuminate, base cuneate, 

acute or narrowly obtuse, sparsely covered with urticating hairs 

or glabrescent, pinnately veined. Inflorescence terminal, consisting of a staminate main axis with a single pistillate basal 

branch; staminate axis racemose, 5-20 cm long, with flowers 

numerous, 1 per node; pistillate branch racemose, 3-10 cm 

long, elongating to 25 cm in fruit, flowers numerous, 1 per 

node; axes sparsely covered with urticating hairs; staminate 

bracts lanceolate, -1 mm long; pistillate bracts lanceolate, 

1.5-2 mm long, apex attenuate. Staminate pedicel 3-5 mm 

long; bud oblong-ellipsoid (or ovoid), 3-4 mm long, apex 

acute; sepals 3, narrowly oblong, 3-4 mm long, apex acute; 

disc segments 8-9, ligulate, 1.5-2.5 mm long, in whorl 

exterior to stamens; stamens 14-17, outer whorl of 8-9 

alternating with and basally fused to disc segments, inner 

cluster of 6-9; filaments slender, 2-3.5 mm long; anthers 

sagitate, -1 mm long. Pistillate pedicel -0.5 mm long, jointed; 

sepals 6, lanceolate, -1.5 mm long; ovary globose, -0.5 mm in 

diameter, very densely covered with urticating hairs; styles 

2.5-3 mm long, connate about one-quarter to one-half of 

length, usually slender, inner surface papillate. Fruiting pedicel 

2-3 mm long; sepals persistant, 3-4 mm long; capsule 

3-lobed, -4.5 x 7 mm, sparsely covered with urticating hairs. 

Seeds ellipsoid-globose, -3.5 mm long, pale brown with darker 

brown spots. 

DISTRIBUTION.-The Guianas and Brazil (Amapa, Maranhao); 20 collections studied, of which 15 are from the Guianas 

(G:2, S:2, FG: 11). 

ECOLOGY.4ccurs in clearings or other disturbed areas of 

moist to wet forest. Flowers and fruits throughout the year. 

SELECTED SPECIMENS ExAMINED.dUyana: Mabura Hill, 

Maas et al. 5910 (MO, NY, U); Mabura Hill, Pipoly & Boyan 

7573 (NY, US). Surinam: Nassau, Lanjouw & Lindeman 2618 

(U); Brownsberg Nature Preserve, Webster 24119 (DAV, GH, 

MO, NY, U). French Guiana: Montagne de Kaw, Feuillet 562 

(CAY, U); Saul, Granville 2783 (CAY, P, U, US); Saul, 

Mori & Gracie 18414 (NY); Montagne de Kaw, Hoff5.548 

(CAY, U); S of airfield at Saul, Skog et al. 7378 (CAY, NY, 

NOTES.-The two collections from Guyana representing the 

westernmost population of the species differ from all other 

collections examined in their broader disc segments, ovoid 

(rather than oblong-ellipsoid) staminate buds, and thicker style 

branches. 

P, US). 

3. Tragia tabulaemontana L.J. Gillespie 

FIGURE 14 

Tragia tabulaemontana L.J. Gillespie, Novon, 4:333, figs. 3,4,6, 8, 10, 1994b. 

[Type: French Guiana, Sommet Tabulaire, about 40 km SE de Saul, 

Granville 3637 (holotype, US!; isotype, CAY!).] 

DESCRIPTION.-Twining vine, monoecious. Stems slender; 

stems and petioles puberulous, sparsely to moderately hirsute, 

and with long-stipitate glandular hairs 0.5-1 mm long and 

urticating hairs 0.2-0.5 mm long. Petiole 0.8-3.2 cm long; 

blade thin-chartaceous, narrowly obovate or narrowly elliptic, 

5-14 x 2-5.5 cm, margin serrate with serration apex obtuse 

and minutely glandular, apex acuminate, base narrowly cordate 
40 SMITHSONIAN CONTRIBUTIONS TO BC ITANY 
NUMBER 86 41 

FIGURE 14 (left).-Tragia tabulaemontana L.J. Gillespie: A, habit; B, leaf 

blade, upper surface; C, upper part of inflorescence showing single open 

staminate flower with staminate buds above; D, staminate flower; E, pistillate 

flower; F, columella and persistent sepals on infructescence axis; G, mericarp of 

dehisced capsule with enclosed seed; H, seed, ventral view (A-E based on 

Granville 3632 US; F based on Granville 3576, CAY; G,H based on Granville 

3632 CAY). 

with sinus 2-6 mm deep, sparsely hirsute on both surfaces, 

venation pinnate, basal secondary vein pair diverging at a more 

acute angle than upper secondary veins. Inflorescence terminal 

on short shoots or appearing leaf-opposed, racemose, 4-9 cm 

long, axes puberulous with numerous long-stipitate glandular 

hairs and scattered short urticating hairs; single pistillate flower 

at basal node, staminate flowers numerous above, 1 per node; 

bracts narrowly triangular-ovate, 1-1.4 mm long. Staminate 

pedicel 1 - 1.7 mm long, puberulous; bud triangular-ovoid, - 1 

mm long, apex obtuse; sepals 3, very broadly ovate, 1-1.2 mm 

long, sparsely hirsute particularly along margin and at apex; 

disc-like structure obtuse-triangular, intrastaminal, adnate to 

base of stamens; stamens 3, filaments 0.5-0.6 mm long, 

conical, slender at apex, highly dilated at base, anthers 0.2-0.3 

mm long. Pistillate pedicel -1 mm long, puberulous, hirsute at 

apex, with glandular hairs along margin; sepals 6, lanceolate, 

-2 mm long, with urticating and glandular hairs; ovary 3-lobed, 

-1 x 1.8 mm, densely covered with urticating and glandular 

hairs; styles 2.5-3.5 mm long, connate about two thirds of 

length, arms recurved. Fruiting pedicel 2-6 mm long; sepals 

persistent, reflexed; capsule 3-lobed, -4.5 x 7-8 mm, sparsely 

covered with urticating hairs and sometimes glandular hairs; 

columella persistent with 3 perpendicular slender apical arms. 

Seeds subglobose, 3.8-4 mm in diameter, pale dull yellow with 

brownish markings. 

DISTRIBUTION.-~OWn only from Sommet Tabulaire in 

French Guiana; two collections studied. 

EcoLoGY .--Occurs in moist submontane forest on the 

southern and western slopes of Sommet Tabulaire, collections 

from 550-600 m. The two collections, both with flowers and 

fruits, were made in August. 

SPECIMENS EXAMINED.-French Guiana: Sommet Tabulaire, -50 km SE de Saul, Granville 3576 (CAY, U, US). 

4. Tragia volubilis L. 

Tragia volubilis L., Sp. pl., 980, 1753. [Type: Jamaica, Herb. Linnaeus 

(holotype, LINN, microfiche IDC 1103.1!).] 

DESCRIPTION.-Twining vine, monoecious. Stems slender, 

becoming somewhat woody; stems and foliage sparsely to 

densely covered with urticating hairs. Petiole 0.5- 1.5 cm long; 

stipules 2-3 mm long; blade chartaceous, narrowly triangularovate, 3-8 x 1-2.5 cm, margin serrate, apex acuminate, base 

truncate or truncate-cordate, 3-veined at base. Inflorescence 

axillary (Guiana specimens), terminal on short shoots or 

appearing leaf-opposed, racemose, 1-4 cm long, axes pubescent; single pistillate flower at basal node, staminate flowers 

numerous above, 1 per node; bracts narrowly ovate-triangular, 

1- 1.5 mm long. Staminate pedicel - 1 mm long, with urticating 

hairs; bud narrowly ellipsoid, 0.8-1 mm long, apex broadly 

acute; sepals 3, ovate, -1 mm long, apex acute, glabrous except 

at base; disc absent; stamens 1-3, filaments 0.2-0.3 mm long 

and wide, anthers -0.3 mm long. Pistillate pedicel -1.5 mm 

long, very densely pubescent; sepals 6, ovate, - 1.5 mm long, 

apex acute, glabrescent or sparsely pubescent; ovary 3-lobed, 

-1 mm in diameter, very densely covered with urticating hairs; 

styles -1 mm long, connate one-half of length, arms divergent. 

Fruiting pedicel 1.5-4 cm long, pubescent; sepals persistant, 

-2 mm long; capsule 3-lobed, 3-3.5 x 5.5-6.5 mm, pale brown 

with dark brown markings, covered with urticating hairs; 

columella persistent, slender with expanded 3-pointed apex. 

Seeds ellipsoid-globose, -3 mm long, pale with reddish brown 

markings. 

DIsTRlBUTI0N.-Widespread in the Neotropics from Guatemala and the West Indies to Bolivia, Argentina, and Brazil, 

apparently absent from the central Amazon Basin, introduced 

in Africa; 50 collections studied, of which 18 are from the 

Guianas (S: 1, FG: 17). 

ECOLOGY.-A weed of open and disturbed areas, found 

along the coast in Surinam and French Guiana. 

SELECTED SPECIMENS EXAMINED.-kham: Bergendane 

Majo, Foche 1027 (U). French Guiana: vicinity of Cayenne, 

Broadway 51 (NY, US); Ile de Cayenne, Prevost 1492 (CAY); 

Cayenne, Rothery 75 (NY). 
Appendix 1 

Numerical List of Accepted Taxa 

1. Omphalea L. 

2. Pera Mutis 

1-1. Omphalea diandra L. 

2- 1. Pera bicolor (Klotzsch) Muell. Arg. 

2-2. Pera decipiens (Muell. Arg.) Muell. Arg. 

2-3. Pera glabrata (Schott) Baill. 

3- I. Dalechampia affinis Muell Arg. 

3-2. Dalechampia attenuistylus Armbruster 

3-3. Dalechampia brevicolumna Armbruster 

3-4. Dalechampia brownsbergensis Webster & Armbruster 

3-5. Dalechampia dioscoreifolia Poepp. 

3-6. Dalechampia fragrans Armbruster 

3-7. Dalechampia heterobractea Armbruster 

3-8. Dalechampia magnoliifolia Muell. Arg. 

3-9. Dalechampia megacalpa Armbruster 

3. Dalechampia L. 

3-10. Dalechampia micrantha Poepp. 

3-1 1. Dalechampia olympiana Kuhlman & Rodrigues 

3- 12. Dalechampia papillistigma Armbruster 

3-13. Dalechampia pawibracteata Lanj. 

3-14. Dalechampia scandens L. 

3- 14a. Dalechampia scandens var. scandens 

3-14b. Dalechampia scandens var. fallax Muell. Arg. 

3-15. Dalechampia stipulacea Muell. Arg. 

3- 16. Dalechampia tiliifolia Lam. 

3- 17. Dalechampia aff. cissifolia Poepp. 

3-18. Dalechampia aff. triphylla Lam. 

4. Haematostemon (Muell. Arg.) Pax & K. HOW. 

4- 1. Haematostemon guianensis Sandwith 

5. Plukenetia L. 

5-1. Plukenetia loretensis Ule 

5-2. Plukenetia polyadenia Muell. Arg. 

5-3. Plukenetia supraglandulosa L. J. Gillespie 

5-4. Plukenetia verrucosa Smith 

5-5. Plukenetia volubilis L. 

6- 1. Tragia fendleri Muell. Arg. 

6-2. Tragia lessertiana (Baill.) Muell. Arg. 

6-3, Tragia tabulaemontana L. J. Gillespie 

6-4. Tragia volubilis L. 

6. Tragia L. 

42 
Appendix 2 

Collections Studied 

(Numbers in parentheses following collection numbers refer to species numbers in taxonomic treatments and in Appendix 1) 

Guyana 

Atkinson, D.J., 114 (4-1) 

Beckett, J.E., 8455 (5-4) 

Beckett & Kortright, 8640 (5-2) 

Boom, B.M., 7 1 17 (2-3) 

Boom & Gopaul, 7326 (2-1) 

Boyan, R., 39 = FD 7863 (2-1) 

Cruz, J.S. de la, 3186 (1-1) 

Davis, T.A.W., 113 = FD 2104 (2-3); 238 = FD 2229 (2-1); 385 = FD 2381 

Ek et al., 1 I87 (3-1 1) 

Fanshawe, D.B., 177 = FD 2913 (2-1); 206 = FD 2942 (2-3); 647 = FD 3383 

Cook, C.D.K., 44 (3-13b) 

(2-3); 40 = FD 2812 (2-1) 

(2-3); 1123 = FD 3859 (4-1); 1212 = FD 3948 (2-3); 1443 = FD 4149 

(2-1); 1993 = FD 4729 (1-1); 2316 = FD 5052 (2-1); 2869 = FD 6016 

(4-1); 2957 = FD 6287 (2-3); 3383 = FD 6947 (5-2) 

FD (= Forest Department of British Guiana), 3923 (3-7); 4434 (3-1); 5409 

Gillespie, L.J., 1955 (3-13b); 4300 (2-1) 

Gillespie & Gopaul, 1009 (3-1); 1993 (3-1) 

Gillespie & Persaud, 1046 (3-13a); 1058 (2-3) 

Gillespie et al., 1654 (3-13a); 1655 (3-13a); 1678 (6-1); 1781 (3-13a); 1832 

Gleason, H.A., 880 (3-1) 

Graham, E.H., 258 (2-1) 

Graham, V., 399 (3-1) 

Harrison, S.G., 714 (3-13a); 1769 (3-13a) 

Hitchcock, A.S., 16770 (3-13a) 

Hohenkerk, L.S., 701 (2-3) 

Irwin, H.S., BG-71 (3-13a); 200 (2-3); 781 (3-1) 

Jansen Jacobs et al., 1751 (2-2); 1828 (3-7) 

Jenman, G.S., 109 (1-1); 495 (1-1); 3617 (2-3); 4088 (3-12); 4138 (2-1); 4718 

Lall, H., 312 (3-13a) 

Maas et al., 5910 (6-2); 7222 (3-13a); 7560 (2-3); 7673 (2-1) 

Maguire & Fanshawe, 23301 (2-1); 32639 (2-1) 

McDowell & Gopaul, 2264 (3-12); 2436 (3-1) 

McDowell et al., 2037 (3-16) 

Pinkus, A.S., 194 (2-1) 

Pipoly, J.J., 7371 (2-3); 7428 (2-3); 10588 (2-1); 11302 (3-1) 

Pipoly & Boyan, (6-2) 

Pipoly & Samuels, 11723 (3-1) 

Pipoly et al., 9705 (2-3) 

Sandwith, N.Y., 425 (2-1); 1565 (3-7) 

Schomburgk, 114 (2-1); 594 (2-3); 610 (3-13a); 783 (3-10); 784 (3-10) 

Schomburgk, Ri., 1070 (2-2); 1071 (2-2) 

Schomburgk, Ro., 580 (2-1); 685 (2-2); 686 (2-2) 

Smith, A.C., 2418 (2-3); 2558 (2-3); 2658 (2-2); 3558 (6-1) 

Tillet & Tillet, 45834 (5-2) 

Tillet et al., 44950 (5-2); 44954 (5-2) 

Wilson-Browne, G., 241 = FD 5714 (2-3); 305 = FD 5734 (2-3) 

(2-3); 5978 (3-15) 

(3-16) 

GUPPY, N.G.L., 203 = FD 7179 (5-1); 627 = FD 7642 (2-2) 

(2-3); 4772 (2-1); 6302 (2-1); 7894 (1-1) 

MMS & Maas, 25 1 1 (2-3) 

Surinam 

Acevedo, P., 6022 (5-3) 

Acevedo et al., 6026 (3-3) 

Armbruster & Herzig, 85-101 (3-6); 85-102 (3-4); 85-103 (3-17); 85-104 (3-6) 

Boldingh, I., 3010 (1-1); 3015 (1-1) 

BW (= Boschwezen, Suriname Forest Service), 240 (3-1); 893 (5-4); 2406 

Daniels & Jonker, 1109 (5-2) 

Florschutz & Maas, 2592 (3-1); 2596 (3-7) 

Foche, H.C., 1027 (6-4) 

Gonggrijp, J.W., 69 (3-1) 

Hekking, W.H.A., 1049 (3-13a) 

Heyde, N.M., 433 (1-1); 459 (3-1) 

Heyde & Lindeman, 50 (1-1); 111 (1-1); 225 (3-7) 

Hostmann & Kappler, 143 (3-1) 

Irwin et al., 54825 (1-1); 54897 (5-5); 55556 (1-1); 55831 (3-13a); 57504 (2-2) 

Kappler, A., 1887 (3-1); 1888 (3-14a); 1889 (3-7); 1913 (3-1) 

LBB (= Lands Bosbeheer), 8565 (5-4); 12534 (J.T. Sterringa) (3-13a); 13687 

(C. Vreden) (3-7) 

Lanjouw, J., 794 (3-7); 867 (1-1) 

Lanjouw & Lindeman, 1045 (1-1); 1114 (3-13a); 1147 (1-1); 1807 (3-13a); 

Lindeman, J.C., 4922 (2-1); 5111 (1-1); 5548 (1-1); 6129 (3-7); 6134 (5-2) 

Lindeman et al., 214 (3-10); 286 (3-1); 636 (5-5); 655 (3-17) 

Maas et al., 2320 (3-6) 

Maguire et al., 54028 (3-1) 

Mennega, A.M.W., 483 (1-1) 

Mori et al., 8693 (3-6) 

Oldenburger et al., 264 (2-2); 3 18 (2-2) 

hlle, A., 152 (3-1); 159 (3-1); 221 (5-4); 273 (1-1) 

Schultz & Donselaar, 10642 (1-1) 

Schulz & Wessels Boer, 10052 (3-1) 

Stahel, G., 82 (2-1); 273 (2-3) 

Stahel & Gonggryp, 41 = BW 3018 (1-1); 190 (1-1) 

Teunissen, P.A., LBB 14503 (1-1) 

Versteeg, G.M., 812 (1-1) 

Webster, G.L., 24119 (6-2); 24138 (1-1); 24143 (3-13a) 

Webster & Armbruster, 24124 (3-4) 

Webster & Mohr, 24122 (3-17) 

Went, F.A.F.C., 548 (1-1) 

Wessels Boer, J.G., 454 (3-10); 958 (3-7); 1108 (3-1) 

(1-1); 4627 (1-1); 4785 (2-1); 4847 (2-3); 5383 (1-1); 6316 (1-1) 

2618 (6-2); 3164 (1-1) 

Schulz, J.P., 7284 (5-2); 8690 (1-1) 

French Guiana 

Alexandre, D.Y., 93 (3-15); 106 (3-13a); 107 (6-4); 112 (3-15); 113 (3-13a); 

227 (3-13a); 377 (6-4) 

Armbruster & Edwards, 95-03 (3-14) 

Barthelemy, D., 145 (3-15) 

Beekman, F., 59 (3-5) 

Benoist, R., 835 (3-13a); 893 (5-4); 1069 (1-1); 1262 (3-13a); 1635 (1-1) 

Billiet & Jadin, 1613 (3-17); 4332 (3-13a) 

Black et al., 55-17589 (1-1) 

Broadway, W.E., 51 (6-4); 444 (3-13a); 631 (3-15); 641 (3-15) 

43 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Cremers, G., 4647 (1-1); 4998 (3-8); 7075 (3-8); 8386 (3-7); 8707 (6-2); 9325 

Cremers & Hoff, 10608 (3-13a) 

Crevaux, (3-13a) 

Croat, T., 74330 (6-2) 

Deward, 240 (2-3) 

Feuillet, C., 417 (3-5); 538 (3-13a); 561 (6-2); 562 (6-2); 570 (6-2); 1754 

Foresta, H. de, 182 (3-8) 

Gamier, 127 (3-15) 

Granville, J.-J. de, 260 (3-15); 265 (3-15); 317 (3-8); T-1047 (1-1); 2770 (1-1); 

(6-2); 9429 (3-15); 9627 (2-3); 9820 (3-1) 

(3-16); 2862 (3-15); 2969 (3-13a); 3015 (3-7) 

2783 (6-2); 3576 (6-3); 3626 (5-3); 3637 (6-3); B.4413 (5-2); B.4846 

(3-7); 5315 (5-3); 5753 (1-1); 6940 (3-13a) 

Granville et al., 7292 (3-5); 10783 (5-3) 

Grenand, 647 (5-2) 

Irwin et al., 48181 (1-1); 48498 (1-1); 48642 (1-1); 48647 (1-1) 

Jacquemin, H., 1664 (1 - 1) 

Larpin, D., 566 (3-7) 

Lemoine, M., 7907 (1-1) 

Hoff, M., 5357 (3-15); 5548 (6-2) 

Maas et al., 2226 (3-5) 

Melinon, E., 305 (6-4); 376 (3-1) 

Merlier, GY203 (6-4); GY245 (6-4) 

Moretti, C., 461 (5-2); 572 (3-7); 698 (3-1) 

Mori & Gracie, 18414 (6-2); 18668 (5-2) 

Mori et al., 14874 (6-2); 15026 (3-15); 23721 (3-14) 

Oldeman, R.A.A., BC-9 (2-3); T-347 (1-1); T-349 (1-1); B-731 (3-1); B-802 

Poncy, O., 4 (3-15); 89 (3-5) 

Prevost, M.F., 649 (2-3); 661 (3-3); 665 (3-3); 1451 (6-4); 1456 (3-13a); 1492 

Raynal-Roques, A., 20006 (3-5) 

Rech, 75 (3-1) 

Rothery, H.C., 75 (6-4) 

Sagot, P.A., 22 (5-4); 511 (1-1); 512 (3-13a); 513 (3-1) 

Service Forestier, 311M (2-1); 3077 (3-13a); 4328 (3-13a); 7909 (1-1); 7969 

Skog et al., 7378 (6-2); 7427 (3-13a) 

Wachenheim, G., 25 (3-13a) 

Mori, S., 15673 (5-2) 

(3-15); T-831 (3-15); 1472 (3-3); 2067 (3-5); B.3227 (3-5) 

(6-4); 1673 (3-3); 1808 (3-15) 

91-1) 
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Webster, G.L., and W.S. Armbruster 

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W.S. Armbruster, and R. Secco, 

editors, Flora of the Venezuelan Guayana. Volume 5. 
Index to Scientific Names 

(Synonyms are in italics) 

Acidoton Sw. 

Apodandra Pax & K. Hoffin. 

Astrococcus Benth. 

loretensis (Ule) Pax & K. Hoffm. 

Bia Klotzsch 

lessertiana (Baill.) Muell. Arg. 

Chaetocarpus Thwaites 

Clutia L. 

Dalechampia L. 

aff. cissifolia Poepp. 

aff. triphylla Lam. 

affinis Muell. Arg. 

amazonica (Ule) Macbride 

attenuistylus Armbruster 

brevicolumna Armbruster 

brevipes Briquet 

brownsbergensis Webster & Armbruster 

cissifolia Poepp. 

dioscoreifolia Poepp. 

fragrans Armbruster 

guianensis Klotzsch 

heterobractea Armbruster 

heterophylla Vahl 

magnoliifolia Muell. Arg. 

megacarpa Armbruster 

micrantha Poepp. 

mollis Vahl 

olympiana 

papillistigma Armbruster 

parvibracteata Lanj. 

peruviana Lam. 

pruriens Griseb. 

roezliana var. amazonica Ule 

rubifomis Spreng. 

ruboides H.B.K. 

scandens L. 

scandens var. scandens 

scandens var. fallax Muell. Arg. 

sidaefolia H.B.K. 

spathulata var. amazonica (Ule) Pax and Hoffmann 

stipulacea Muell. Arg. 

tiliifolia Lam. 

triphylla Lam. 

Elaeophora Ducke 

abutifolia Ducke 

polyadenia (Muell. Arg.) Ducke 

Euphorbia L. 

Haematostemon (Muell. Arg.) Pax & K. Hoffi. 

coriaceus (Baill.) Pax & K. Hoffm. 

guianensis Sandwith 

Megalostylis Spencer Moore 

poeppigii Spencer Moore 

Omphalea L. 

diandra L. 

megacarpa Hemsley 

Pera Mutis 

bicolor (Klotzsch) Muell. Arg. 

decipiens (Muell. Arg.) Muell. Arg. 

distichophylla (Mart.) Baill. 

ferruginea (Schott) Muell. Arg. 

glabrata (Schott) Baill. 

nitida (Benth.) Jabl. 

tomentosa (Benth.) Muell. Arg. 

bicolor Klotzsch 

bicolor var. nitidum Benth. 

bicolor var. tornentosum Benth. 

decipiens Muell. Arg. 

ferrugineum Schon 

glabratum Schott 

schomburgkianum Benth. 

caribeus Urb. 

abutifolia (Ducke) Pax & K. Hoffm. 

integrifolia Vahl 

loretensis Ule 

macrostyla Ule 

peruviana Muell. Arg. 

polyadenia Muell. Arg. 

supraglandulosa L.J. Gillespie 

venucosa Smith 

volubilis L. 

Peridium 

Phyllanthus L. 

Plukenetia L. 

Pogonophora Miers ex Benth 

Tragia L. 

fendleri Muell. Arg. 

japurensis Muell. Arg. 

lessertiana (Baill.) Muell. Arg. 

tabulaemontana L.J. Gillespie 

volubilis L. 

47 
baaka tiki 

baboenoot 

baboenotto 

brongwirie 

chichinat 

graine de I’anse 

hachiballi 

Index to Vernacular Names 

hatsiballi 

hatsiballi koelnoe 

idaballie 

liane papaye 

mekoekwaire 

ouabe 

kamua-tan 

palaramauloeloe 

peprehoedoe 

pirikraipjo 

sooke 

stijo 

UIT 

48 

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