SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 28 

A Monograph of the Genus Eperua 

(Leguminosae: Caesalpinioideae) 

Richard S. Cowan 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1975 
ABSTRACT 

Cowan, Richard S. A Monograph of the Genus Eperua (Leguminosae: Caesalpinioideae). Smithsonian Contributions to Botany, number 28, 45 pages, 13 

figures, 2 tables, 1975.-AIl available data are brought together in a monographic 

treatment of the legume genus Eperua. Eighteen taxa are considered of which 

four are described as new: E. duckeana, E. obtusata, E. grandiflora ssp. guyanensis, and E. jenmanii ssp. sandwithii. Also one new combination is made: E. 

glabriflora (Ducke) Cowan. In addition to gross morphology, anatomy of leaf 

epidermis and palynology of most of the species are presented for the first time. 

Pollen morphology is particularly instructive with respect to the classification 

derived principally from vegetative and floral morphology. 

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

in the Institution’s annual report, Smithsonian Year. SI PRESS NUMBER 5383. SERIES COVER DESIGN: 

Leaf clearing from the katsura tree Cercidiphyllurn japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data. 

Cowan, Richard S. 

A monograph of the genus Epeura (Leguminosae: Caesalpinioideae) 

(Smithsonian contributions to botany, no. 28) 

Bibliography: p. 

Supt. of Docs. no.: SI 1.29:28 

1. Eperua. I. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to botany, 

no’. 28. 

QK1.542747 no. 28 [QK495.L52j 581’.08s [583’.323] 75-619031 
Contents 

Introduction ................................................ 

Nomenclatural History of the Genus ........................ 

Gross Morphology ............................................. 

Epidermal Anatomy ............................................... 

Pollen and Pollination ............................................ 

Ecology and Phenology ............................................ 

Geography ............... .................................. 

Economics ....................................................... 

Phylogenetic Speculations .......................................... 

Eperua Aublet .................................................... 

Key to the Taxa of Eperua .................................... 

1 . E . schomburgkiana .......................................... 

3 . E . rubzgznosa ............................................... 

a . E.r. var . rubiginosa ................................... 

b . E.Y. var . grandifiora .................................. 

4 . E . falcata .................................................. 

5 . E . venosa .................................................. 

6 . E . leucanthn ............................................... 

7 . E . duckeana .............................................. 

8 . E . glabriflora .............................................. 

9 . E . bzpiga ................................................. 

10 . E . obtusata ................................. 

11 . E . grnncliflora .......................................... 

a . E.g. ssp . guyanensis ................................. 

2 . E . glabra .................................................. .. 

* . 

b . E.g . ssp . grandiflora .................................. 

b . E.j. ssp . sandwithii ................................ 

12 . E . jenmanii ............................................... 

a . E.j. ssp . jenmanii ................................... 

13 . E . pwpzirea ............................................... 

14 . E . oleifern ............................................... 

a . E.o. var . oleifera ..................................... 

b . E.o. var . campestris ................................. 

Appendix 1 : Excluded Names ................................ 

Appendix 2: Lists ................................................. 

Literature Cited .................... ......................... 

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111 

A Monograph of the Genus Eperua 

(Leguminosae: Caesalpinioideae) 

Richard S. Cowan 

Introduction 

During an expedition to the upper Orinoco 

Basin in 1950-51, I first saw living material of 

the genus Eperua and was impressed by the 

beauty of the flowers and the anomalous pseudotubular corolla. In the years that followed I had 

occasion to work with various species of the group 

and developed a deep interest in becoming better 

acquainted with the entire genus. Then, in working on a floristic treatment of the caesalpinioid legumes for the Flora Vene zuela, it was necessary to 

revise the genus in order to be able to treat the 

species of Epertia authoritatively. Presented herein 

is a summary of everything I have been able to discover about the group from the field, the herbarium and from the literature. 

Numerous persons should be acknowledged as 

having been helpful to the completion of this 

monograph, but I especially want to call attention 

to Nancy Goor, who prepared the drawings, to 

John Wurdack, who read the manuscript, to Joan 

Nowicke for her assistance and encouragement 

with the palynology section, to E. S. Ayensu for 

his advice in making and interpreting anatomical 

preparations, and particularly to the Smithsonian 

Research Foundation, which generously provided 

grant assistance. 

Nomenclatural History of the Genus 

generic names, Eperua and Parivoa, in widely separated parts of his ground-breaking Histoire des 

plantes de la Guiane francaise (1775). Even though 

the greatly elongate, pendent inflorescences and 

falcate leaflets of E. falcata Aublet appear quite 

dissimilar to the short, erect inflorescences and 

nonfalcate leaflets of E. grandiflora (Aublet) Bentham, Schreber (1791) recognized the two species 

to be congeneric. Following the custom of the 

time (one of which he was especially observant) he 

coined a new generic name, Dimorpha, to cover 

the two dissimilar species. 

Willdenow (1799) treated both species in his 

edition of the Species Plantarum, providing a new 

name, Panzera, for Eperua but retaining Schreber’s 

name Dimorpha for the other species. At the same 

time he added a second species, D. tomentosa 

(Aublet) Willdenow, based on Pariuoa tomentosa 

Aublet, which is usually assigned to Crudia. De 

Candolle (1825) was the last of the major encyclopedists to treat this assemblage as two genera, 

Eperua and Parivoa, for Bentham and Hooker in 

their Genera Plantarum (1865) combined the two 

concepts under the one name Eperua, admitting 

only the two original Aublet species. Since then, 

various authors have described one or two species, mostly in connection with floristic reportsBentham in the Flora Brasiliensis (1870), Miquel 

in Stirpes surinamenses selectae (1 850), Sandwith 

in a review of the genus in Guyana (1931), and 

As so often happened with Aublet genera, the 

first two species were described under different 

Cowan on miscellaneous species in the Guiana 

Highland (1957 and 1958). Only one attempt at 

an overall survey of the species has been made-by Richard S. Cowan, Department of Botany, Rational Museum 

of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. Flora Brasiliensis (1870). 

=ucke in 1940-since Bentham’s review in the 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 1 .-Scanning electron photomicrographs of epidermal morphology: a, pubescence of 

E. rubigimsa, X 855; b, lower epidermis of E. purpurea, X 300 (Maguire 29486, US); c, the 

same, X 1500. 
NUMBER 28 3 

Gross Morphology 

Although differing greatly in stature, all the 

species are trees, some very small (E. oleifera var. 

campestris), but others are among the largest in 

the Amazonian rain forest (E. purpurea). The 

trunks of some of the species are fluted and/or 

buttressed, but the presence or absence of buttresses 

may be related to the substrate, rather than being 

an obligate genetic characteristic. The bark, insofar as there are any data, is smooth except for rather 

obvious corky lenticels, and gray to brownish-gray 

or gray-black. 

Pubescence is generally simple and microscopic 

(X 15 or more) to macroscopic but never very 

long, except on the androecium. In E. rubiginosa 

the hairlets are compound, each one branching 

irregularly in a dendroid fashion (Figure la); 

those of E. schomburgkiana are microscopically 

bifid, with the appearance of two hairlets arising 

from the same point. 

The stipules are often foliaceous, up to several 

centimeters long and wide, free from each other, 

and situated at the two sides of the petiole base. 

In species characterized by such stipules, however, 

they may fail to develop completely and then appear as small, inconspicuous “flaps”; thus stipules 

are less valuable as a source of key characters than 

would be the case if they were always well expressed. Several species have the stipules partly to 

completely united laterally into a single bifid or 

entire body in the leaf axil. 

The leaves are uniformly even-pinnate, but the 

number of pairs of leaflets is not so constant. Four 

species regularly have only bijugate leaflets (E. 

bijuga, E. glabriflora, E. obtusata, and E. uenosa) 

but most have 3- to 5-jugate ones, sometimes having some of the leaves with only two pairs. There 

is always a well-developed cylindrical petiole and 

rachis, as well as obvious petiolules; the blades of 

the leaflets are usually equilateral and straight, 

but in three species, all with long, pendant inflorescences, the leaflets are obviously falcate, the two 

halves of each leaflet consequently more or less 

unequal. Pubescence on the leaves is rare, but the 

undersurface of the leaflets in E. pzirpuyea is obscured by a microscopic layer of tangled filaments 

of wax (Figure lb,c). Venation of the leafllets is 

generally inconspicuous, sometimes even obscure, 

but a strong marginal vein is present in E. pzirpurea and E. oleifera var. campestris; sporadically, 

submarginal veins occur in other species but not 

so regularly that they provide a means of recognizing the group. The leaflets of most taxa have pellucid punctae, and sometimes they are also present in flower parts, especially in the sepals but 

never in the petal. In most instances, because of 

the thickness of the parts, it is necessary to view 

the leaflets or flower part against a strong light 

source in order to see the punctae, but in E. oleifera and E. pzirpurea, they are more obvious. 

The inflorescences are basically the same in all 

the species, although two species groups can easily 

be distinguished on inflorescence type alone. 

Whether elongate-pendent (up to several meters 

long) or short-erect, a central axis bears a number 

of racemosely disposed lateral branches, each of 

which is a raceme. Thus, the differences between 

the two types of inflorescence are essentially the 

result of one developmental characteristic-the 

much greater elongation of the peduncle and the 

axis intervals between branches in the elongatependent type. A broad bract, often caducous to 

deciduous, subtends each flower, the pedicel of 

which bears, at various levels, a pair of more or 

less opposite, caducous to persistent bracteoles, very 

similar in form, size, and pubescence to the bracts. 

The cupular hypanthium is somewhat inequilateral in a few taxa (E. bijuga, E. grandflora, E. 

jenmanii) but always lined on the inner surface 

with a palisadelike series of nectaries that project 

above the rim of the hypanthium or are included 

within it. Vogel (1968), comparing Eperua to 

Hymenaea, considered that each flower has a single nectary which in the former is greatly increased 

in surface area by radial cleavage or invagination 

(“radiale Zerkliiftung”). My observations lead me 

to interpret each of Vogel’s “folds” as one of several nectaries. In either case, the hypanthium is 

tightly filled around the gynophore with the nectariferous tissue. 

Four strongly imbricate sepals become free from 

each other at the rim of the hypanthium-they 

presumably form the outer layer of the hypanthium. Those portions of the outer surface of each 

sepal, not partially covered by the adjacent ones, 

bear the pubescence (if any) characteristic of the 

taxon; the overlapped portions are always glabrous, 

thinner, often scarious. The sepals are unequal to 

varying extents in several species, the outer pair, or 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FILLRE 2.-Morphology of Eperua species. E. purpurea (Maguire et a1 60112, F): a, habit, X y2; 

B, stipular body, X 1. E. jenniaiiii var. jenninnii (Sandwith 710, IVY): c, foliaceous stipules, ca. 

X 1. E. rubiginosa (Irwin et a1 57579, NY): d, bud showing teiminal extrahypanthial nectaries, 

ca. X 1. E. jennzanii (drawn from plate in protologue): e, essential organs. E. purpicrea (Maguire 

et a1 60112, F); j, open, partly dissected flower, X 1.5; g, expanded petal, ca. X 1. E piirpurea 

(Ll. IVilliams 13990, US): h, fruit, X '/2. 
XUMBER 28 

at least the dorsal one, larger and more strongly 

concave/cucullate. In three species (E. glabriflora, 

E. obtzisata, and E. rubiginosa) the outer two sepals have a single, large, raised gland near the apex 

on the outer surface. In E. rubiginosa at least these 

appear to be extrahypanthial nectaries that secrete 

actively just prior to anthesis (see section on Pollen and Pollination). The sepals vary from green 

to red to whitish, partly in response to aging of the 

flowers, and they generally are thickish. The 

corolla consists of four minute petalodia (up to 10 

mm long but usually 0.5-5 mm long) and one 

large to relatively very large glabrous petal that 

envelops the essential organs of the flower, producing the appearance of a tubular corolla such as is 

characteristic of the Bignoniaceae. The range of 

colors-white, red, purple, lilac, pink, etc.-is also 

like many of this family, so it is not surprising that 

there have been family misidentifications by otherwise highly competent field botanists! The width 

of the petal is often greater than the length so that 

there is a strong overlap of the lateral margins 

in bud. In most species the petal is oblate or flabelliform, sometimes tapering obcuneately in a clawlike base, but more often the petals are truncate 

basally. In only one species (E. venosa) does the 

petal bear any pubescence, and in this instance it 

is only slightly strigulose externally at the extreme 

base. In bud the petal is strongly crumpled around 

the infolded stamens and coiled style. 

The androecium consists of ten stamens, of 

which as many as five may be reduced in size and 

in some species sterile. In most species the smaller 

anthers appear to produce normal pollen, although 

its viability is not known. The one dorsal stamen 

is free from the tube formed by the connation of 

the other filaments, or it may be briefly joined at 

the extreme base; it appears to be fertile in all 

species, but its filament is usually shorter and may 

be somewhat different in vestiture. The filament 

tube is equilateral-the union of the filaments to 

about the same level-or strongly inequilateral by 

the progressively greater union of the filaments 

toward the ventral side of the flower (opposite the 

insertion of the petal). Where there is dimorphism 

in proportions, the length of the filaments is always 

greater when they bear the larger anthers characteristic of each species, those bearing smaller or 

sterile anthers being about the length of the 

free dorsal one that is invariably fertile. The longest pubescence observed in the genus occurs on the 

filament tube of several species. The anthers are 

versatile, dorsifixed, oval to narrowly oblong, 

almost invariably glabrous, and only rarely with 

apiculate connectives. The thecae are sometimes 

conspicuously but minutely tessellate and always 

opening before anthesis. 

Within the base of the filament tube the apices 

of the nectaries are usually clearly evident. According to Vogel (1968), the nectar collects in the 

space at the base of the petal, but this seems very 

unlikely in view of the posture of the flower and 

the adaxial insertion of the petal, unless the nectar 

is very viscous. On the other hand, it appears entirely reasonable to imagine that the nectar collects 

within the filament tube, which serves as a reservoirlike structure, ensuring an abundance of 

pollinator-attracting nectar at anthesis (see section 

on Pollen and Pollination). If this interpretation 

is correct, then the deeper, zygomorphic filament 

tube of some taxa must be a more effective reservoir, at least in more or less horizontally oriented 

flowers. 

The gynoecium is the least variable part of 

the flower, consisting of an obtuse to capitate 

stigma, an elongate style (four to ten times the 

length of the ovary), an oblong-obovate to oblongoblanceolate ovary, and a short gynophore (generally 

shorter than the ovary). The stigma is usually 

entire, but in two species (E. leucantha and E. 

venosa) it is somewhat bilobed. The style is always 

placed at the dorsal margin of the broad ovary 

apex; that is, the dorsal margin of the ovary continues, scarcely with any change in alignment, as 

the dorsal surface of the style. The gynophore typically is borne in the center of the regular hypanthium cup but more or less eccentrically in those that 

are zygomorphic. The ovary contains several ovules. 

The fruits are always woody, dehiscent (sometimes violently so in a plant press!), flat (except 

for dorsal winglike expansions perpendicular to 

the pod surface in E. glabyiflora and E. purptirea), 

and more or less oblong to subquadrate. Of those 

that have the oblong form, several species have 

falcate, arcuate, or scimitar shapes. One or two 

oval, flat seeds are found in each fruit; the seed 

coat is thin-chartaceous and dark colored. 

Seedlings of only two species are known certainly, and the first leaves of these have fewer pairs 

of leaflets than is typical of mature foliage. Seeds 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

apparently germinate on the surface of the soil, 

the cotyledons emerge from the testa (“phanerocotyly”) sending an elongate epicotyl “canopyward” and an equally elongate primary root into 

the substrate. The hypocotyl is very short, only a 

few millimeters long with no special morphology 

to delimit it precisely. 

Epidermal Anatomy 

In an effort to clarify further the phylogenetic 

relationships of the taxa comprising this genus, 

the anatomy of the epidermis was studied. The 

technique of applying a liquid plastic to the surface of a leaflet and then observing the resulting 

“peel” microscopically, failed to yield useful data. 

Likewise, scanning electron microscope techniques 

for observing the cellular details were unsuccessful. 

I am indebted to E. S. Ayensu for his assistance in 

evolving the following method for rapidly assaying 

the range of variation in structure of the epidermis 

of thick, coriaceous tropical leaves. 

Small bits of dried leaf tissue roughly 5 mm in 

length and breadth were placed directly into a 

mixture of equal parts of 10 percent nitric acid 

and 10 percent chromic acid and left for at least 

24 hours or until the bits became translucent. The 

samples were then thoroughly washed using a jet 

of water from a plastic washing bottle. This treatment separates the upper and lower epidermis 

from the mesophyll, most of which is washed away, 

but any remaining nonepidermal tissues can be 

removed with a fine camel-hair brush in a watch 

glass of water under a dissecting microscope. Small 

fragments of the epidermis are stained for about 

five minutes in Toluidine Blue, washed, and then 

mounted in a water-miscible mountant. The result 

is only a temporary slide, prepared in a very few 

minutes, which can be studied at magnifications 

up to and including oil immersion. If permanent 

slides are indicated by the preliminary examination, other bits of the cleared sample can be run 

through conventional stain/dehydration series for 

mounting in one of the permanent mountants. 

The Toluidine Blue stain “bleeds” into the temporary mountant, but otherwise slides prepared by 

this method are usable for at least a few weeks. 

In Eperua the upper epidermis lacks stomata, 

so all the data reported below are based on the 

observations of the lower epidermis. There is more 

uniformity than anticipated in epidermal characters, for all eighteen taxa showed paracytic-type 

stomata which, for the most part, are evenly distributed over the surface; in a few taxa, the stomata occur in clusters. (I realize this is a greatly 

simplified technical description of stomata relative 

to the ontogenetic terminology proposed by FrynsClaessens and van Cotthem (1973), but applicaFIGURE 3.-Lower epidermis of species of Eperua, X 400: a, E. oleifera var. campestris (Schultes 

10337, US); 6, E. schornburgkiana (de la Cruz 2172, NY); c, E. purpurea (Maguire 29486, US). 
NUhlBER 28 7 

tion of this sort of approach to tropical trees is, at 

least, difficult!) Figure 3a-c indicates the range of 

variation in size of stomata and of the nonstomatal cells. Also, the extremes of convoluting of 

the cell walls is illustrated by this figure as well; 

in other taxa the cell walls are intermediate between these extremes. There appear to be no phylogenetic conclusions to be deduced from epidermal 

characters in this genus, for there is no correlation 

with the phylogenetic arrangement suggested by 

gross morphology and by palynology. 

The lower epidermis of E. purpurea requires 

Taxa 

separate consideration, because it is so different 

from that of all the other taxa. Figure Ib (SEM, 

X 300) reveals that the glaucous appearance of 

the leaflet surfaces is due to a layer of waxlike 

material, and Figure Ic (SEM, X 1500) demonstrates that the material occurs in the form of 

branching threads. Doughnut-shaped stomata are 

easily distinguished from the “ping-pong-ball” 

shaped bodies which I have referred to as “wax 

glands.” The drawing in Figure 3c indicates that 

these are produced on ordinary epidermal cells 

but not on guard cells or subsidiary cells. 

TABLE I.-Summary of pollen data (under “size,” figures outside parentheses = 

range of one-half or more of measurements; figures in parentheses = minimum 

Source of pollen Thickness km 

Size Sexine Nexine Aperture Ektexine morphology 

1. E. schomburgkiana 

3. E. rubiginosa var. 

rubiginosa 

4. E. jalcata 

5. E. venosa 

6. E. leucantha 

7 E. duckeana 

8 E. gla bra flora 

9 E. bijuga 

11. E. grandaflora ssp 

grandijlora 

12. E. jenmanat ssp. 

jen man1 i 

13. E. purpurea 

14 E. oleafera var. 

oleifera 

Persaud 166 (US)-Guyana (71-)74-78 2-3 

de la Cruz 2172 (NY)-Guyana (74-)78-84* 1.5-2 

0.6-1 3-colporate 

1-1.5 

punctate-rugulose 

Lanjouw 1209 (US)-Suriname (82-)91-97 4-5 

(-101) 

Froes 26649 (US)-Brazil 78-88’ 4 

Pires 48876 (US)-Brazil (104-)116- 5-6.5 

123(-126)’ 

Tillett 45791 (NY)-Guyana (91-)104- 4-5 

114(-123) 

Maguire 53597 (US)-Venezuela (88-)94-104 2-2.5 

Ducke 23289 (US)-Brazil (88-)95-101 2.5-3 

(-1 12) 

(-104) 

Froes 21 104 (NY)-Brazil (91-)96-104 2-2.5 

(-1 10) 

Ducke 244 (iYY)-Brazil (65-)69-71(-75) 2-2.5 

Ducke 14A (F)-Brazil (55-)59-62(-65) 2-2.5 

INPA 6065 (US)-Brazil (62-)66(-70)’ 2.5 

Ducke 20312 (US)-Brazil (65-)70-74(-78) 2-2.5 

Froes 32988 (US)-Brazil (74-)80(-88)’ 2.5 

Pires 50673 (US)-Brazil (60-)65-72(-78) 2 

Pires 51659 (F)-Brazil (45-)47-53(-60) 1.5-2 

Maguire 33136 (US)-Venezuela (58-)68-71(-77) 1.5 

Gleason 837 (US)-Guyana (62-)65(-69)’ 1.5-2 

Maguire 41922 (US)-Venezuela 65-71(-73) 4-5 

Prance 15513 (US)-Brazil 65-71(-74) 5 

Ducke 235a (9Y)-Brazil 48-61(-65)’ 4-5.5 

Ducke 213 (NY)-Brazil (51)55-58(-66)’ 4.5-5.5 

1.5-2 3-colporate 

1-1.5 

1.5 3-colporate 

0.7-1.5 

0.7-1 3.porate 

1.5 3-porate 

1-1.5 

1 3-colporate 

1.5 3-colporate 

1-1.5 

1-1.5 3-colporate 

1.5 

0.7-1 3-colporate 

0.7-1 

1 3-colporate 

1-1.5 

1.5 3-colporate 

1.5 

0.7-1 3-colporate 

1-1.5 

verrucose 

reticulate 

(heterobrochate) 

puncta te-rugulose 

punctate-rugulose 

punctate-rugulose 

punctate 

punctate 

punctate 

punctate 

reticulate 

(homobrochate) 

reticulate 

(homobrochate) 

*Based on less than 25 measurements. 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 4.-Pollen of Eperua species. E. schomburgkiana (Persaud 166 US): a, X 1000; b, X 3000. 

E. rubiginosa (Lanjouw 1209, US): c, X 850; d, X 3000. 
in only 

(1 968), 

In this 

NUhfBER 28 

Pollen and Pollination 

Anthesis has been observed and detailed 

one species, E. schornburgkiana, by Vogel 

who described it as a bat-Dollinated Dlant. L I 

species anthesis begins in early evening, but this 

may not be true for other taxa, even in those whose 

flowers are pollinated by bats. In all species the 

sepals open partially, the bases remain imbricate, 

and the petal which is strongly crumpled in bud 

expands. At the same time the infolded filaments 

reflex and assume the posture characteristic of the 

taxon involved. In E. schornburgkiana a faint odor 

of crushed leaves (“ein Krautiger Geruch”) is evident in the flower at this stage (Vogel, 1968). 

Pollen is already released before anthesis, and, 

at least in some buds, I have observed numerous 

pollen grains on the stigma which before anthesis 

is more or less enclosed by the coiled style. The 

latter characteristic has often been interpreted as 

an adaptation that at least partially prevents selfing, but differential ripening of the anthers and 

the stigma is surely a more effective barrier where 

selfing is deleterious. After anthesis and pollination, all the flower parts except the gynoecium are 

shed; only in E. rubiginosa does the filament tube 

remain intact around the carpophore of the developing fruit. 

There is considerable variation in the form, size, 

and ornamentation of the pollen grains among the 

species of this genus; Table 1 is presented in combination with scanning electron micrographs to 

summarize the variability in a concise way. Observations of overall size and wall thickness were 

made by light microscopy at X 400, based on 25 or 

more grains on one or a series of slides made from 

a single flower. Only grains were measured which 

appeared to be mature, intact, not damaged by 

developmental accidents or by preparation of the 

slides; otherwise the choice of grains to measure 

was random, Two different collections, often from 

two distinct geographic locations, were used for 

each of the twelve species examined. Pollen was 

not available from two species, E. glabra and E. 

obtusata, because of the paucity of material. Fasbender (1959) described the pollen grains of nine 

of the species, but her observations were limited to 

those possible with light microscopy, and some of 

the fine details revealed by the SEM are consequently absent from this important pioneering 

paper. 

9 

Because the grains are flattened equatorially, the 

material, especially in slide preparations, tends to 

be oriented in polar view. For this reason, the 

measurements given in Table 1 represent the 

height of the triangle, i.e., the distance from any 

one apex to the base of the triangle. Using this parameter, the grain size ranges from 45 pm to 126 pm. 

The pollen of one species, E. falcata, falls into 

Erdtman’s (1952) “very large spores” class, measuring in excess of 100 pm, three others just short 

of this size (E. rubiginosa, E. venosa, and E. leucantha), and, in fact, only two samples had grains 

less than 50 pm. In general, the largest grains are 

associated with species judged to be advanced 

sporophytically. 

The thickness of the sexine and nexine was measured midway between two apertures and ranges 

from 1.5 to 6.5 pm for the sexine and 0.7 to 2 pm 

in the case of the nexine. Although the nexine 

was of relatively uniform thickness within each 

grain, the thickness of the sexine was more variable. 

In some species the sexine thickens near the apertures (E. glabriflora, E. bijuga, and E. falcata). 

The other taxa have pollen with the sexine 

equally thick throughout, or it actually thins toward the apertures (E. schornburgkiana, E. uenosa, E. purpurea, E. oleifera). What significance 

evolutionarily or biologically to attach to these observations is, at least, uncertain. 

COMPREHEKSIVE DESCRIPTION OF THE POLLEN.- 

Grains single, triangular in polar view with an 

aperture at each angle, flattened equatorially, 3- 

colporate, rarely 3-porate, the ektexine more or 

less punctate, punctate-rugulose, verrucose, or 

reticulate. 

Most of the taxa examined (ten of twelve species) 

had pollen with colporate apertures and the colpi 

short or reduced, but in two closely related species, E. uenosa and E. leucantha (Figures 5, 6), the 

apertures are porate and probably represent the 

ultimate reduction of the colpus. The presence of 

occasional porate apertures in the “colporate” 

species appears to reinforce this view. In another 

species pair, E. purpurea and E. oleifera, the apertures are distinctive due to the width of the colpus 

(Figure 9). 

Based on morphology, two very generall groupings can be recognized-unspecialized pollen and 

specialized pollen. “Unspecialized pollen grains,” 

3-colporate with the ektexine punctate or punctate- 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 5.-Pollen of Eperua species. E. faEcata (Pires et a1 48876, US): a, X 850; b, X 3000. 

E. venom (Maguire et a1 53597, US): c, X 750; d, X 3000. 
NUMBER 28 

FIGURE B.-Pollen of Eperua species. E. leucantha (Ducke 23289, US): a, X 850; b, X 1400. 

E. duckeana (Ducke 23734, US): c, X 1400; d, X 3000. 

11 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 7.-Pollen of Eperua species. E. glabriflora (Ducke 14a, F): a, X 1400: b, X 3000. 

E. bijuga (Ducke 20312, US): c, X 1000; d, X 3000. 
NUMBER 28 

FIGURE 8.-Pollen of Efierua species. E. grandifEora (Pires 50673, US): a, X 1000; b, X 3000. 

E. jenrnanii (Maguire 33136, US): c, X 1000; d, X 3000. 

13 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 9.-Pollen of Eperua species. E. purpurea (Maguire 41922, US): a, X 1350; b, X 3000. 

E. oleifera (Ducke 235a, NY): c, X 1700; d, X 3000. 
NUMBER 28 15 

rugulose, are found in six species: E. glabriflora, 

E. bijuga, E. jenmanii, E. grandiflora, E. duckeana, 

and E. schomburgkiana (Figures 4a,b; 6c,d; 7, 8); 

the latter two could have been separated by the 

rugulose character of their ektexine, but the difference is not sharply defined. The category “Specialized pollen grains” includes those types that appear 

to be derived or advanced in some characteristic: E. 

oleifera and E. purpurea (Figure 9) have grains 

with the ektexine homobrochate-reticulate and, 

moreover, they have the most prominent colpi; E. 

leucantha and E. uenosa (Figures 5, 6) are the 

only taxa with porate apertures, regarded as advanced (see discussion above), but have a relatively 

unspecialized ektexine; E. rubiginosa (Figure 4) 

has an ektexine that is conspicuously verrucose, a 

condition that could be derived by an elaboration 

of the larger elements or folds in a punctaterugulose ektexine, such as is found in E. venosa 

and E. leucantha (Figures 5, 6); the pollen grains 

of E. falcata (Figure 6) are distinctive by virtue of 

the reticulate ektexine in which, unlike the reticulum of E. purpurea and E. oleifera pollen, the 

brochi are of very variable sizes (heterobrochate). 

There have been very few observations of the 

visitors to flowers of any species of Eperua and 

none that give unequivocal evidence of the true 

pollinators. H. S. Irwin (pers. comm.), however, 

has observed large black ants and hummingbirds 

visiting flowers of E. rubiginosa during daylight 

hours, bats at dawn and at dusk. Which of these 

groups effect pollination is unknown at this time 

and all may be involved, the ants in selfing, the 

bats and hummingbirds in outcrossing. In E. rubiginosa, however, ants may very well be pollinators; in a photograph generously provided by 

Irwin of an inflorescence branch, one can clearly 

see a drop of liquid (“a sticky drop,” fide Irwin) 

at the apex of each bud just before anthesis. This 

is one of the species with the two outer sepals bearing a single large dorsal “gland” near the apex. I 

suspect that these are extrahypanthial nectaries 

that function just before anthesis to “advertise” 

to potential pollinators the coming attractions of 

the hypanthial nectar flow. In other words, pollinators would be attracted to the mature buds SO 

that large numbers would be present at anthesis. 

Since ants are not likely to move from tree to tree 

of the same taxon, it would seem that ant pollination, if it does occur regularly, is tantamount to 

describing the plants as self-compatible. While all 

this may be true, it is speculation at the moment 

and needs field checking to verify. At least in some 

other tropical legumes the same swarms of ants 

are present in the inflorescences at anthesis, but 

Mary K. Arroyo (pers. comm.) believes that in 

these instances the ants serve to protect the tender, 

often succulent flower parts against predators. This 

view has also been expressed by Thomas Elias and 

Howard Irwin at the New York Botanical Garden 

(pers. comm.). The situation may be exactly the 

same in Eperua, but at this moment there are no 

published observations to solve the riddle. 

One may as well credit as pollinators the bats 

and hummingbirds that visit the flowers of this 

and closely related species. The long-exserted 

stamens, surpassed considerably by the elongate 

style, appear to provide for the shedding of pollen 

on the backs of such visitors as they enter the flower 

in search of nectar. Vogel (1968) considered E. 

schomburgkiana, E. leucantha, E. falcata, and E. 

rubiginosa to be chiropterophilous, largely because 

of the morphology of the flower and inflorescence, 

rather than because he or others had observed bat 

visitors to the flowers covered with pollen. He 

pointed out that the following are melittophilous 

species: E. bijuga, E. purpurea, E. jenmanii, E. 

oleifera, and E. grandiflora, but he also apparently 

reached that conclusion on the basis of morphology, 

rather than on field observations. 

If, in fact, Eperua species “cover all their bets” 

by employing both selfing and outcrossing in accomplishing fertilization, this would seem to have 

considerable adaptive advantage. Such a situation 

is of course intermediate between the position 

taken by Fedorov (1966), Baker (1959), and others 

(that low density of individuals and asynchronous 

flowering among tropical trees naturally leads to 

self-compatibility and selfing) and the views of 

Bawa (1974), Ashton (1969), and others who hold 

either that outcrossing is the rule or that it is the 

more commonly effective breeding system. One of 

the more rewarding areas for investigation in biology presently is that concerned with these and 

related phenomena. 

Ecology and Phenology 

There is always a dearth of ecological data when 

one is studying tropical plants, especially when one 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

must rely almost exclusively on collection data. As 

Brainard once remarked in his work on Viola in 

North America, one suspects that distribution maps 

reflect more the itineraries of collectors than the 

range of the plant species they purport to show. In 

the neotropics, one is faced by this same phenomenon, intensified by the difficulty of reaching much 

of the region, even now when technology is opening the interior regions to exploration. Add to this 

the fact that in the rain-forest regions where so 

many of the tree legumes abound in numbers of 

species and individuals, there is a several-monthlong rainy season that discourages most collectors. 

So one is left with compiling data from collections 

made primarily during the drier parts of the year 

in accessible areas. The following, admittedly incomplete, account is based solely on data from 

labels on collections, mostly made under these 

conditions. 

Plants of the genus Eperua are almost invariably 

lowland rain forest trees, often along rivers and 

streams (this latter condition may reflect the fact 

that most collecting has been done along rivers, 

the principal transportation routes where roads 

have not yet penetrated). Some species are best 

developed in that part of the forest above the influence of annual flooding, while others occupy the 

regularly flooded portions (especially E. falcata). 

Very often Eperua species grow along blackwater 

streams, and Janzen (1974) in a review of existing 

data on the cause of such rivers and lakes cites 

leaching of “humic acids and probably other toxic 

organic compounds” from Eperua leaves, among 

others. He would also contend that the evergreen 

leaves of most species is an adaptation to poor soils, 

too poor to permit replacement of leaves regularly. 

And many Eperzia species do occur on white or 

brown sands in forest or savanna scrub. Two taxa 

occur in drier, nonforest situations: E. purpurea 

generally is a plant of savanna scrub, and E. oleifera var. campestris occurs in grassy savannas. 

Dominance of one Eperua species or another is so 

pronounced in some areas as to approach the kind 

of forest one finds in temperate zones dominated 

by one or a few tree species. Janzen concluded that 

seeds of Eperun probably “will be found to be 

very toxic.” This could explain the relatively pure 

stands of wallaba species-a result of the development of a built-in predator deterrent, perhaps 

even a biocide, in the seeds. Certainly the large 

number of seedlings sometimes seen simultaneously germinating beneath a parent tree suggests 

at least “unpalatable” seeds. He discarded the possibility of mast-fruiting as a means for the genus to 

overcome predator destruction. Other factors involved in the evolution of tropical forest were reviewed by Ashton (1969). 

Soils in the hylaea are often sandy and in Guyana 

there is a regular relationship between the kind of 

soil and the Eperua species it supports-white sand 

and brown sand soils usually have different species. 

According to Ducke (1940), E. glabrifiora is 

found in places with moist acidic soil of white sand 

and black humus, E. schomburgkiana on the marshy 

banks of small forest rivers and streams. 

Altitudinally the range for the genus is not great, 

all taxa occurring below 1000 m elevation: E. bijuga 

occurs along watercourses at the mouth of the 

Amazon River, near sea level, and most species 

occur below 500 m; Eperua jenmanii has been collected up to 1000 m and E. falcata at 700 m elevation. In this respect, Eperua taxa are typical of the 

vast majority of woody legumes in the neotropicsthey are essentially a lowland component, albeit a 

very critical one in terms of biomass, of the tropical rain forest. 

Fanshawe’s classic paper on the vegetation of 

Guyana (1952) is particularly instructive with respect to Eperua ecology. Although the area considered there (Guyana) is not typical of all parts of 

the range of Eperua, it is certainly a good representation of much of it. Using Beard’s classification 

of climax vegetation in tropical America (1944), 

Fanshawe described the components of 28 formations, in at least seven of which one or more species 

of Eperzia occurs. Some species are narrowly restricted, and none occur in either his Coastal Zone 

or his Southern Upland Region, but one species, 

E. falcatn, is able to tolerate or to take advantage 

of a wide range of soil types, water supply, elevation, and to endure man-made or natural disturbances, such as clear felling and burning for shifting 

agriculture or storm destruction. In most areas, 

however, it typically forms a nearly pure stand on 

the floodplain along the rivers. Just above its range, 

on the land behind the floodplain and above 

annual flooding, E. rubiginosa is dominant (H. S. 

Irwin, pers. comm.). In the Lowland Region, 

which stretches from the northwest, southeasterly 

to the southern uplands, forests of Eperua in com- 
NUMBER 28 17 

bination with various other subdominants and 

associates cover large areas. These occur in two 

forms principally and on two general soil types: 

(1) Dry Evergreen Forest on the central white 

sand peneplain or on redistributed white sands 

derived from it; and (2) in Swamp Forest on alluvial flats alongside rivers. The same sort of pattern prevails in the mountainous sandstone region 

of the Pakaraima Mountains. 

From the incomplete data available it appears 

that most species flower during the (relatively) 

dry season at its driest, others at the end of the dry 

and the beginning of the rainy season; fruits usually mature in the rainy periods. Eperua bijuga, 

however, appears to flower and fruit during the 

rainy season (April to August); E. leucantha and 

E. falcata flower and fruit during the dry season 

(September to March). 

Geography 

The species of Eperua are unusually restricted 

for a lowland legume genus. All known species occupy the northeastern quarter of the South American continent from loo south to the north coast 

and from about 73” west longitude, east to the 

Atlantic, boundaries that coincide well with those 

of the Amazonian Hylaea. Of the eighteen taxa 

recognized in the genus, only five occur south of 

the equator and only four are unknown in Venezuela and the Guianas. There are large inexplicable gaps in the distribution where considerable collecting should have turned up one or more taxa. 

The whole Brazilian mid-Amazon and mid-Rio 

Negro areas have had relatively good attention 

by collectors, but the map shows an absence of 

Eperua localities. The vast preponderance of both 

individuals and kinds occur in the southeastern 

Colombian-southwestern Venezuela-Guianas Arc. 

A secondary concentration of taxa occurs in the 

Manaus area. Collecting in the distribution gaps 

listed above, as well as in the species-rich lowland 

forest area of southeastern Colombia may be expected to yield many additional distribution records and very likely undescribed taxa. 

Economics 

The name “wallaba” is applied to timbers of 

several species of Eperua, the commonest, E. falcata, is generally called “soft wallaba.” The wood 

of E. jenmanii, usually called “ituri wallaba,” is 

indistinguishable commercially (but is different 

even to the unaided eye) from soft wallaba (Aitken, 1930). The trees grow rather rapidly and regeneration is relatively rapid, about thirty years 

from cutting to cutting. Wallaba forest grows 

naturally on agriculturally useless white sands, 

which are capable of producing an average of 600 

cubic feet of marketable wallaba timber per acre 

(Aitken, 1930). 

The sapwood is dingy white with resinous 

streaks, but the darker heartwood is decidedly 

resinous. The heavy, hardwood splits rather cleanly, 

although the gummy exudate makes working it 

difficult in the same way as does “fat pine” wood. 

A1 though paper-pulp experiments indicate it to 

be an uneconomical source of pulpwood, the timbers are used widely throughout Guyana for house 

framing, vat staves, fence poles, telephone poles, 

and shingles, as well as for firewood and charcoal. 

It is particularly resistant to decay in contact with 

water and mud. (During a visit in 1954 I was told 

that a seaplane ramp, being built of wallaba 

planks into one of the northern Guyana rivers, 

should be serviceable for at least 25-30 years, even 

though submerged!) 

Ducke (1940) reports that the two forms of E. 

oleifera, E. bijuga, and E. purpurea had been in 

cultivation for four to six years in 1940 in the 

Jardim Botanic0 do Rio de Janeiro; E. falcata is 

planted in Trinidad, West Java, and Cameroon; 

and E. jenmanii has been grown in Mexico, Singapore, Cameroon, Jamaica, and Trinidad. 

Phylogenetic Speculations 

There seems to be little cause to doubt that 

Eperua, Heterostemon, Elizabetha, and Paloue 

form a close-knit cluster of genera, even though 

they have been considered (Engler & Prantl, 1894) 

to be more distantly related by virtue of the fact 

that the latter three genera, and others, have three 

to five petals and/or their flower buds are enclosed 

in enlarged bracteoles before anthesis. But it is 

even more speculative to suggest where this genus 

cluster should be associated, Vogel (1968) believed 

that Eperua and Hymenaea are closely related, a 

view shared by Lee and Langenheim (1975) in 

their monograph of the latter genus. However, I 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

am unable to see any greater relationship between 

them than Eperua appears to have with many 

other nearby genera, and resolution of such questions must be postponed pending completion of a 

survey of generic characters in progress. 

Eperua is readily divisible on the form of the 

inflorescence into subgeneric categories, but recognition of sections or subgenera in such a relatively 

small genus, while tempting, really serves no practical or scientific purpose. Moreover, there is also 

the possibility that the long, pendent type of inflorescence may have evolved more than once, 

although it seems unlikely; at least there is no 

necessity for assuming that that development has 

occurred. At the subspecific level I have recognized 

both varieties and subspecies; the latter I view as 

incipient species with substantial geographic discontinuity, as well as some morphological difference (s) which are either of insufficient importance 

or regularity to characterize a species. 

The accompanying diagram (Figure 10) of possible relationships has all the weaknesses of such 

attempts to illustrate phylogeny in one plane; however, it is constructed on the following assumptions: 

1. Leaves with many pairs of leaflets are less advanced than those with fewer pairs. 

2. Intrapetiolar, nonfoliaceous stipules are a refinement in contrast with lateral, foliaceous ones. 

3. Long pendent inflorescences are a relatively 

advanced trait. 

4. All stamens fertile are less advanced; progressive reduition in size of anthers, increased atrophy 

in them, and possible loss of viability of the pollen 

in the reduced anthers occur in more advanced 

taxa. 

TABLE 2.-Characters and characteristics of Eperua species, used to derive the 

presumed phylogenetic relationships shown in Figure 10 

Character Characteristic 

A. Inflorescence length 

and posture 

short, erect (0) 

elongate, pendent (1) 

short, regular (0) 

and form long, zygomorphic (1) 

10 fertile (0) 

less than 10 fertile (1) 

00000011111111 

00010000011111 

B. Stamina1 tube length 

C. Stamen number fertile 

D. Sepal nectary absent (0) 

present (1) 

equal size/form (0) 

unequal size/form (1) 

lateral (0) 

intrapetiolar (1) 

foliaceous (0) 

nonfoliaceous (1) 

zygomorphic (1) 

11111111001111 

00011100000011 

00011101000011 

00000000101100 

E. Sepal form 

F. Stipule position 

G. Stipule form 

H. Hypanthium form regular (0) 

00001100000000 

00001100000000 

I. Stigma form obtuse to capitate (0) 

J. Pollen aperture colporate (0) 

bilobed-capitate (1) 

porate (1) 

K. Pollen ektexine 0011 punctate or punctatereticulate or verrucose (1) 

001100000 rugulose (0) 
NUMBER 28 

5. Zygomorphic hypanthium and staminal tube 

are relatively advanced. 

6. Unequal sepals with extrahypanthial nectaries are evidence of advancement. 

7. Bilobed stigmas are an elaboration of simple 

capitate, capitellate, or truncate stigmas. 

8. Punctate-psilate ektexine is interpreted as less 

advanced than punctate-rugulose, verrucose, and 

reticulate, in that order. 

9. Colporate pollen apertures preceded porate 

ones evolutionarily. 

Some of these trends are characteristic of the subfamily Caesalpinioideae generally, especially the 

tendency for reduction in leaflets per leaf, and the 

progressive sterilization of some members of the 

androecium. The presence of only a single petal 

in Eperua is only one example of the oft-expressed 

tendency for reduction in petal number in the 

subfamily. 

As W. H. Wagner has often remarked in discussing his “ground plan” method of representing 

phylogenies in this graphic form, one of its great 

strengths is that it forces one to study very intently 

all the characters of a group, to evaluate them 

and to judge how they interrelate. In Table 2, only 

19 

the most significant characters are included, together with their character-states, for each species. 

Subspecific taxa are not treated separately in the 

table because all parts of each species are the same 

with respect to the characters/characteristics included. 

The immediate precursors of extant Eperuaspecies must have had leaves with numerous pairs 

of leaflets, foliaceous-lateral stipules, erect inflorescences bearing flowers whose sepals were of equal 

size and form and lacking extrahypanthial nectaries, with a regular cupular hypanthium, a short 

regular staminal tube bearing ten functional anthers enclosing pollen with punctate ektexine and 

germinating by means of three colporate apertures 

on nonbilobed stigmas. The solid circle in Figure 

10 represents such an assemblage of unspecialized 

characteristics, from which the modern species 

have evolved by acquisition of more advanced 

traits. Eperua schomburgkiana, for example, is advanced beyond the ancestral state principally by 

its sepals that are unequal in size and form: on 

the other hand, E. falcata is relatively far removed 

by its accumulation of six character-states which 

I have interpreted to reflect advancement. The 

- fa I c a t ~(ACEFGK) 

FIGURE 10.-Ground-plan diagram of phylogenetic relationships in Eperua. 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

concentric semicircular lines represent evolutionary stages, each with a progressively larger number of specializations. The capital letters along the 

radiating lines, as well as those labeling the unnamed hypothetical taxa and in the formulas in 

parentheses after each specific epithet connote the 

characters listed in Table 2. 

Many speculations can be made on the basis of 

this diagram at once but these are probably 

obvious. What would be most interesting would 

be the discovery in the field, or herbarium, of some 

of the hypothetical taxa depicted! Such an event 

would tend to validate what is otherwise highly 

putative. 

Eperua Aublet 

Eperzia Aublet, P1. Guian. 1:369, t. 142, 1775. 

Pariuoa Aublet, P1. Guian. 2:756, t. 303, 1775. 

Dimorpha Schreber, Gen. P1. 2493, May 1791. 

Panzera Willdenow, Sp. P1. 2540, March 1799. 

Small to very large trees (5-70 m tall); stipules 

2, often free, foliaceous and conspicuous but sometimes joined in a bifid or entire, compound intrapetiolar body; leaves alternate, paripinnate, the 

leaflets 2- to 6-jugate, usually glabrous, pellucidpunctate, falcate or straight; inflorescence racemes, 

usually compound, long and pendent or short and 

erect, the flower subtended by a small bract at the 

pedicel base and two bracteoles arising at various 

levels on the pedicel; flowers pedicellate, the 

hypanthium cupular, regular to zygomorphic, 

the inner wall obscured by a palisade of nectarproducing glands; sepals four, equal in size and 

shape or the dorsal one larger and more concave, 

the two outer ones sometimes with a single gland 

near the apex dorsally, strongly imbricate; petal 

one, usually glabrous, oblate or flabelliform, obcuneate or truncate basally; stamens 10, all fertile 

or up to 5 reduced to staminodia, the dorsal one 

fertile, free or joined at the base with the tube 

formed by the union of the filaments in their lower 

part; stigma obtuse to capitate, entire or bilobed, 

the style filiform, the ovary oblong-oblanceolate 

to obovate, the gynophore centric to nearly so in 

the hypanthium; fruits woody, dehiscent, scimitarshaped to subquadrate, the seeds one to few, oval, 

flattened laterally, 

TYPE-SPECIEs.--EPerua fukata Aublet, PI. Guian. 

1:369, t. 142, 1775. 

Key to the Taxa of Eperua 

1. Inflorescence long-pendent, to 2.5 m long; ovary usually longer than gynophore ....... 2 

Inflorescences erect or at least not pendent, only a few centimeters long; ovary and gynophore about equal in length ........................................................................ .7 

2. Leaflets falcate; stipules intrapetiolar, bifid or entire ............................................... 3 

Leaflets not falcate; stipules free from each other, lateral at the base of the petiole ...... 5 

3. Leaflets mostly bijugate and 14.5-22 cm long, 8-9 cm wide, the base regularly inequilateral, 

the venation prominent; petal very shortly unguiculate, sericeous externally at the base .. 

................................................................................................................... 5. E. venosa 

Leaflets mostly 3-jugate, not as in the preceding; petal truncate basally, sessile, glabrous. . 4 

4. Petal white, 25-30 mm long, 40-50 mm wide; buds with a gray bloom (minutely puberulous); inflorescences 6-9 dm long, the axis minutely puberulous, the axis of the lateral 

branches mostly 10-15 mm long at anthesis; staminal tube strigose-sericeous; fruits 

minutely strigulose ................................................................................ 6. E. Zeucmttha 

Petal pink to scarlet, 13-18 mm long and 19-25 mm wide; buds brownish-strigulose; inflorescences 10-14 cm long, the axis somewhat microstrigulose, the axis of the lateral 

branches about 30-50 mm long or even longer at anthesis; staminal tube villose; fruits 

more or less minutely puberulous ............................................................... .4. E. falcata 

5. Plant with many parts minutely tomentulose, the hairs multicellular, irregularly branching, 

rusty-colored (at least X 60), petal dark pink to red or purple-red, 13-20 mm long, 22-35 

mm wide; filaments 35-55 mm long, the anthers 5.5-7 mm long ........ (3. E. rubiginosa) 6 

Plant glabrous in all its parts; petal cream-colored, 28-30 mm long, 35-50 mm wide; filaments about 78 mm long, the anthers 9 mm long .................................. 2. E. glabra 

6. Leaf rachis usually 12.5 cm long or shorter, the leaflets 3-jugate; sepals 13-20 mm long, 

Leaf rachis usually more than 15 cm long, the leaflets 4- or 5-jugate; sepals 21-35 mm 

long, 7-15 mm wide ........................................................... 3b. E. rubiginosa var. grandiflora 

4-9 mm wide .................................................................. 3. E. rubiginosa var. rubiginosa 
NUMBER 28 21 

7. Leaflets bijugate ............................................................................................................. 8 

Leaflets 3- to 6-jugate ................................................................................................................. 10 

8. Apex of leaflets rounded-obtuse, partly retuse ............................................ 10. E. obtusata 

Apex of leaflets acute to acuminate ..................................................................................... 9 

9. Leaflets narrowly revolute marginally; inflorescence axes glabrous; bracteoles 9-10 mm long, 

oval-ovate; pedicels aud sepals glabrous; northwestern Brazil ................... 8. E. glabriflora 

Leaflets plane; inflorescence axes puberulous; bracteoles 5 mm long, oblate; pedicels and 

sepals puberulous; Lower Amazon Valley, especially near mouth of Amazon River, and 

in French Guiana ............................................................................. 9. E. bijuga 

10. Leaflet surfaces strongly discolorous, densely papillate-ceriferous on the undersurfaces (at 

least X 30), the “wax bodies” contiguous, obscuring epidermis; bracteoles obviously 

scarious marginally, persistent at least to anthesis, triangular-ovate ...... 13. E. purfiurea 

Leaflets never ceriferous; bracteoles not as above .............................................. ..11 

11. Petal white; fruits more or less oblong, more than twice as long as wide .......... 12 

Petal pink, lilac, rose-purple, or purple; fruits more or less quadrate, less than twice as long 

as wide ................................................................................................................. 13 

12. Leaflets mostly 5- or 6-jugate; sepals 22-30 mm long; petal 25-35 mm long, 35-60 mm wide; 

gynoecium glabrous; staminal tube equilateral, 10-15 mm long; Guyana and Suriname ...... 

Leaflets mostly 4-jugate; sepals 12-18 mm long; petal 14-25 mm long, 22-35 mm wide; ovary 

and gynophore tomentulose densely; staminal tube somewhat zygomorphic, 3.5-9 mm 

long; vicinity of Manaus, Brazil .................................................. 7. E. duckerma 

13. Petal dark laveuder to purple with darker purple and white markings, mottled, basally 

white, 5-10 cm long and 4-8.5 cm wide .............................................. (12. E. jenmanii) 14 

Petal pink to purple, 2.5-4 cm long aud 2.5-5 cm wide ............................... .15 

14. Bracts, bracteoles, inflorescence axes, pedicels, and hypanthium minutely puberulous; bracts 

4-5 mm long .................................................................. 12a. E. jenmanii ssp. jenmanii 

Bracts and bracteoles glabrous except somewhat ciliolate; inflorescence axes, pedicels, and 

hypanthium glabrous; bracts 2-3 mm long ...................... 12b. E. jenmanii ssp. sandwithii 

15. Petal lilac to purple; branchlets, petioles and rachis microscopically pubescent sparingly; 

hypanthium equilateral; staminal tube villose; southern Amazon Basin ................. 

Petal pink; branchlets, petioles, and rachis glabrous; hypanthium inequilateral; staminal 

tube puberulous; northeast Brazil and Guyana ..................................... 17 

16. Large forest tree with oval to elliptic leaflets, the base rounded-obtuse; petal pale lilac ..... 

Small savanna tree with ovate; cordate leaflets; petal rose-purple or purple ............ 

............................................................................................................ 1. E. schomburgkiana 

........................................................................................................ (14. E. oleifera) 16 

...................................................................... 14a. E. oleifera var. oleifera 

...................................................................... .14b. E. oleifera var. campestris 

ovary and gynophore densely pilosulose; central to south-central Guyana 

.......................................................................... lla. E. grandiflora ssp. guyanensis 

coastal French Guiana and southern Suriname ............. 1 lb. E. grandiflora ssp. grandiflora 

17. Leaflets minutely venulose, the margins revolute; anthers more or less minutely puberulous; 

.... ...... 

Leaflets plane, the venation subobscure; anthers and gynoecium glabrous; northeast Brazil, 

1. Eperua schomburgkiana Bentham 

FIGURES 4, 11 

Epenca scho~nburgkiana Bentham, F1. Bras. 15 (2):226, 1870. 

Large shrub or tree 5-36 m tall, the branchlets 

glabrous, nitid, the stipules 15-43 mm long, 8-22 

mm wide, foliose, venulose, coriaceous, deciduous, 

inequilaterally arcuate-ovate, acute; petioles (1 3-) 

18-25 (-30) mm long, glabrous, the rachis 13-27 

cm long, glabrous; leaflets 5- or &jugate, the petiolules 6-10 (-12) mm long, glabrous, the blades 

sometimes irregularly pellucid-punctate, coriaceous 

to rigid-coriaceous, plane, glabrous, often nitid, 

(4.5-)7-12 (-15) cm long, (2.5-)3.5-5.5 cm wide, 

all lanceolate or the basal one or two pairs lanceolate with the other pairs elliptic, the base of the 

lower one to four pairs obtuse to subobtuse, the 

base of the upper pairs acute, the apex acuminate 

to long-acuminate, often mucronulate, the venation more or less prominulous; inflorescence a terminal raceme of racemes 5-14 cm long, the lateral 

ones in same plane, the axes tomentose, the hairs 

rusty colored and at least some bifid minutely, the 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

bracts and bracteoles caducous, sometimes deciduous, 4.5-5.5 mm long and 4-5 mm wide, broadly 

ovate or oval, concave, tomentose externally; pedicels 11-27 (-35) mm long, tomentose, the hypanthium cupular 4-7 mm long, 5-10 mm in diameter, 

tomentulose externally, the nectaries projecting 

above the hypanthial rim; sepals 22-30 mm long, 

6-1 5 mm wide, elliptic to oblong-elliptic, tomentulose externally, the two outer ones larger, concave; petal white, glabrous, oblate to oblateobovate, 25-37 mm long, 35-60 mm wide; all 

stamens apparently fertile, the filaments 45-60 mm 

long, 9 united in an equilateral tube 10-15 mm 

long, the tube more or less pilosulose or tomentulose, the anthers oval, glabrous, 4.0-5.5 mm long, 

1.5-2.5 mm wide; gynoecium glabrous or nearly 

so, the stigma capitate, the style 45 mm long, the 

ovary 11-15 mm long, 3.5-5 mm wide, rarely a few 

hairlets on the dorsal margin near the base, oblanceolate, more or less arcuate, the gynophore 9- 

17 mm long, glabrous; fruits oblong-oblanceolate, 

25 cm long, 8 cm wide, somewhat nitid, glabrous, 

the carpophore 2 cm long, glabrous; mature seeds 

not seen. 

TYPE CoLLEcTIoN.-Robt. Schomburgk 517 (lectotype K, isolectotypes BM, G, K, P, W), “Essequibo & Rupununy. British Guiana, 1838.” 

DIsTRIBuTI0N.-Riverine forest on sandy soils 

in northern Guyana and in adjacent Suriname. 

Flowering June to October. 

GUYANA: Above Arrawarri mouth, west bank of lower 

Essequibo R., 20 March 1909, Anderson 182 (K, RB); Mazaruni R., no date, A@un 359 (K); upper Mazaruni R., 22 

Sept-6 Oct 1922, de la Cruz 2091 (F, GH, NY, US), 2172 (F, 

GH, NY, US); Mazaruni Station, 10 March 1934, Davis 394 

(FD 2390) (K), 7 March 1941, Fanshawe 644 (FD 3380) (K, 

NY); Tumatumari, Potaro R., 4-6 July 1921, Gleason 332 

(GH, SY, US), 394 (NY) and 412 (NY); Penal Settlement, 3-9 

Dec 1919, Hitchcock 17089 (GH, US); Mazaruni R., Sept 1880, 

Jenrnan 627 (K), June 1889, 5257 (BM, K, NY): Essequibo R., 

Sept-Oct 1881, Jenrnan 1336 (K); no locality, June 1924, 

Persaud 14 (F, NY); Seeba, Oct 1924, Persaud 166 (F, NY, 

us); Essequibo R., Moraballi Creek near Bartica, 28 Aug 

1929, Sandwith 142 (K, NY, RE, U, US); mouth of Moraballi Creek, Essequibo R., 11 Aug 1960, Watson 58 (BM). 

SURINAME: Corantijn, near mouth of New River, 1 Sept 

1935, Rornbouts 170 (NY, U); Wanatoba Falls, Corantijn R., 

26 June 1936, Schulz 7708 (NY, U, US); Corantijn R., 

25 July 1950, Suriname Forestry Bureau 806 (K, IJ). One collection from Brazil [Para, west bank of Rio Pacaja, 15 Oct 

1965, Prance et al 1652 (F, NY, U, US)] is in fruit and, except for its origin far out of the geographic limits of the 

species, represents it rather well. 

LOCAL NAMES AND UsEs.-Ituri wallaba (Persaud 

166); mahaica-bally (Persaud 14); water wallaba 

(Persaud 166, Schulz 7708). 

D~scuss~o~.--Although not very much less advanced than some of the related species, E. schomOurgkiana combines features (foliaceous lateral 

stipules, relatively large number of leaflets, erect 

inflorescences, regular hypanthium and stamina1 

tube, and ten fertile stamens) that seem to locate 

it near the base of a putative phylogenetic scheme. 

It is most nearly related, in any interpretation, to 

E. duckeana which was identified by Ducke as a 

“local form” of E. schomburgkiana. The present 

species differs in having more pairs of leaflets, 

larger flowers, and a glabrous ovary, as well as in 

its distribution. Palynologically, the two species 

are quite similar, the punctate-rugulose ektexine 

persisting through this evolutionary line to such 

advanced forms as E. leucantha and E. venosa, 

which have long-pendent inflorescences and 3- 

porate pollen, rather than the short-erect inflorescences and 3-colporate pollen of this basic stock. 

This species seems a logical candidate for the 

taxon from which those species with long-pendent 

inflorescences may have evolved. Of these, E. rubiginosa may be the least advanced; E. obtusata is 

possibly derived from the same stock, for at least 

both taxa have a conspicuous dorsal gland near 

the apex of each of the outer two sepals, but E. 

obtusata has short, erect inflorescences and other 

relatively less advanced characteristics which relate 

it closely to E. schombwgkiana. 

2. Eperua glabra Cowan 

FIGURE 11 

Eperua glabra Cowan, Brittonia 83231, 9 Jan 1957. 

Tree 5-20 m tall and to 30 cm in diameter, 

the bark variegated colored, conspicuously corkylenticillate, the branchlets glabrous, nitid; stipules 

caducous, foliaceous, falcate-lanceolate, glabrous, 

about 2.5 cm long and 1.5 cm wide; leaves glabrous, 

the petioles (10-)25-35 mm long, the rachis 11.5- 

19 cm long; leaflets 3- or 4-jugate, the petiolules 

5-8 mm long, the blades epunctate, chartaceous, 
NUMBER 28 23 

the lowest pair ovate, cordulate basally, 9.5-14.5 

cm long, 5-7 cm wide, those of the other pairs 

elliptic, 11-17 cm long, 5-7.5 cm wide, the second 

pair obtuse basally, the upper two pairs acute at 

the base, all leaflets long-acuminate abruptly, the 

upper and lower surfaces of the leaflets strongly discolorous, the costa plane above, salient beneath, 

the venules minutely prominulous; inflorescence 

a pendent, terminal raceme of racemes to 3 m long, 

the lateral axes (2-)5-6 cm long, the axes glabrous, the bracts and bracteoles deciduous, glabrous 

except ciliolate marginally, about 2-3 mm long 

and wide, broadly ovate, the bracteoles arising 

near the pedicel base; flowers glabrous, the pedicels about 25 mm long, the hypanthium 5 mm long, 

funnel-form, the nectaries included within the hypanthial cup; sepals 20-22.5 mm long, 8-12.5 mm 

wide, elliptic, acute, two outer ones cucullate; 

petal glabrous, 28-30 mm long, 35-50 mm wide, 

flabelliform, truncate basally, mostly cream-colored 

but greenish at the base and rose-colored marginally, the petalodia ca. 1.2 mm long, oval, obtuse, 

glabrous; androecium glabrous, the stamina1 tube 

subequilateral, 2.5-3.5 mm long, all stamens fertile, 

the filaments ca. 78 mm long, the anthers 9 mm 

long, 2 mm wide, narrowly oblong; gynoecium 

glabrous, the stigma capitate, the style 60 mm 

long, the ovary 12.5 mm long, 3.5 mm wide, 

oblanceolate-oblong, 5-ovulate, the gynophore 8.5 

mm long; immature fruit narrowly oblance-oblong, 

22-30 cm long, 5-9.5 cm wide, arcuate, the carpophores 3.5-4 cm long. 

TYPE COLLECTION.-N.G.L. Guppy 430 (FD 

No. 7445) (holotype NY, isotypes K, NY, US), 

near Chodikar Landing, Acarai Mts., 250 m elevation, Guyana, 17 October 1952. 

cality, the mountainous frontier separating Guyana 

and Brazil, where it is locally abundant. 

DIsTRIBUT1oN.-Known only from the type 10- 

GUYANA: Shodikar Creek, 8-22 Jan 1938, A. C. Smith 2882 

(A, F, G, NY, U, US). 

DIscmsIoN.-The closest relative of E. glabra 

appears to be E. rubiginosa, which it resembles in 

inflorescence morphology, lateral stipules, nonfalcate leaflets, and in having all ten stamens fertile. It differs in being glabrous in most parts, petal 

color (cream-colored) and size, stamen dimensions, 

and in its eglandular sepals. 

3. Eperua rubiginosa Miquel 

FICURB 4, 11 

Eperua rubiginosa Miquel, Stirp. Sur. Select 12, 1850. 

Small to mediumsized tree (infrequently larger) 

5-20 (-30) m tall, the trunk 12-35 (-90) cm in diameter, the bark brownish-gray, papillate-lenticellate, 

the branchlets glabrous, often nitid; stipules 

glabrous, deciduous to persistent, triangular to 

ovate-triangular, 1.5-3 mm long, 1.5-3.5 mrn wide, 

or foliaceous, venulose, broadly oval, 20 mm long 

and 17 mm wide; leaves glabrous, the petioles 

8-20(-30) mm long, the rachis 5.5-22.5 cm long; 

leaflets 3- to 5-jugate (seedlings 2-jugate), the petiolules 4-7 (-9) mm long, the blades epunctate, 

coriaceous to rigidly thick-coriaceous, the margins 

plane to narrowly revolute, 6.5-13 (-23) cm long, 

(3-)4.5-6 (-10.5) cm wide, all elliptic or more commonly the basal pair ovate to lanceolate, the base 

equilateral, obtuse to rounded-obtuse or the second 

and third pairs subobtuse to acute, sometimes attenuate, the apex acute to long-acuminate, mucronulate to mucronate, the venules subobscure to 

subprominulous; inflorescence a pendent, terminal 

raceme of racemes 100-200 (-250) cm long, the 

lateral racemes (2.5-)4-8 (-16) cm long, the axes 

microtomentulose, the hairlets multicellular, irregularly branching, rusty colored, the bracts and 

bracteoles caducous, microtomentulose densely externally, sparingly so on inner surfaces, the bracts 

ca. 2 mm long and 1.5 mIn wide, the bracteoles 2.5 

mm long, 3 mm wide, oval-ovate or oblong-oval, 

concave, arising about one-third of the length of 

the pedicel from its base; pedicels 20-30 mm long, 

1-3 mm in diameter, rusty tomentulose minutely, 

the hypanthium 3.5-5(-10) mm long, 4.5-6 mm 

in diameter, microtomentulose externally; sepals 

green, the two outer ones larger, cucullate and 

with one raised gland externally near the tip, microtomentulose externally, glabrous on the inner 

surface, narrowly oblong to oblong-elliptic to oval, 

fleshy, 13-30 mm long, 4-15 mm wide; petal glabrous, dark pink to red to purple-red, 13-35 mm 

long, 22-50 mm wide, broadly oblate to flabelliform, truncate basally, the four petalodia 0.6-1.2 

mm long, triangular-ovate, glabrous; stamens persistent with young fruits, most or all apparently 

fertile, 9 or 10 united basally in an equilateral tube 

1.5-3 mm long, glabrous except in the tube 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sinuses, 35-70 mm long, the dorsal one shorter, 

united with the tube at the extreme base or even 

higher, the anthers narrowly oblong, brown, 5.5- 

11 mm long, 1.5-2 mm wide, glabrous; stigma capitate, the style 20-80 mm long, glabrous, the ovary 

7.5-10 mm long, 2.5-3.5 mm wide, oblanceolateoblong, truncate apically, densely tomentulose, the 

gynophore 6-1 2 mm long, tomentulose minutely; 

mature fruits to 52 cm long and 12.5 cm wide, oblanceolate, more or less scimitar-formed, arcuate, 

rusty tomentulose. 

3a. Eperua rubiginosa Miquel var. rubiginosa 

FIGURES 2d, 11 

Stipules triangular to ovate-triangular, 1.5-3 mm 

long; leaf rachis 5.5-12.5 cm long, the leaflets 3- or 

4-jugate; inflorescences 10-25 dm long, the pedicels 

about 2 mm or less in diameter just above the 

bracteoles; sepals 13-20 mm long, 4-9 mm wide. 

TYPE COLLECTIOS.-F. W. Hostmann 229 (lectotype U, isolectotype K), Suriname R., Suriname. 

Choice of lectotype is necessitated by Miquel's citation of two syntypes, and this collection was referred to as the type by Amshoff in the Flora of 

Szil-iname. 

D1sTRIBuTIoN.-Frequent to common riverine 

tree, usually above the level of annual flooding in 

central Guyana and Suriname on alluvial deposits, 

coastal French Guiana and northeastern Brazil 

(Amapi). Flowering from July through November; fruiting August to October. 

GUYANA: Arawari Creek, Essequibo R., 22 March 1909, 

Anderson 183 (K); Essequibo and Mazaruni R., Appun 294 

(K); Mazaruni Station, 19 Nov 1934, Davis 404 (FD 2400) (K); 

no specific locality or date, Fanshawe 2104 (NY, U, US); 

swamp forest behind Mission, 22 Sept 1952, Guppy 198 (FD 

7174) (K, NY, U); Kurupukari, Essequibo R., 27 July 1920, 

Hohenkerk 122-B (K); Mazaruni R., Sept 1880, Jenman 768 

(K) and 786 (K); Essequibo R., Sept-Oct 1881, Jenman 1147 

(K); south\vest of Bartica, 3 Dec 1923, Linder I32 (GH, NY); 

Kassikaityu R., upper Essequibo R., 19 Nov 1935, Myers 5672 

(K); no locality or date, Robt. Schomburgk 515 (BM, F, G, 

GH, K, NY, IV), Rich, Schonzburgk 173 (Bhl), 17f (BM, G ,K, 

P), and 181 (P); near mouth of Onoro Creek on Essequibo 

R., 15-24 Dec 1937, A. C. Smith 2741 (A, F, G, NY, U, US). 

SURIK'AME: Onobissi, Coppename R., 6 Nov 1916, Gonggrijp 2577 (6); Trakoekakreek, 21 Oct 1921, Gonggrijp 5515 

(Uj; Kayser .4irstrip, 45 km above junction of Zuid R. with 

the Lucie R., 25 Sept 1963, Irwin et a1 57579 (M, NY, U, US); 

Suriname R., Oct 1844, Kappler 1689 (syntype of E. rubiginosa 

Miquel) (G, P, U, W); Arakukakreek, Kegel 872 (NY) Coppename R., near Raleigh Falls, 10 Sept 1933, Lanjouw 738 

(U); Suriname R., near Kabelstation, 10 Nov 1933, Lanjouw 

I209 (U, US); bank of Marowijn R., south of Albina, 19 Sept 

1948, Lanjouw 5 Lindeman 324 (K, NY, U); Coppename R., 

near mouth of Wayombo R., 23 July 1954, Lindeman 6344 

(U); near junction of Zuid and Lucie Rivers, 7 July 1963, 

Maguire et al 54014 (F, G, GH, NY, RB, U, US); Raleighvallen, Coppename R., 24 Sept 1954, Mennega 102 (U); 

Corantijn R., near junction with New R., 1 Sept 1935, Rombouts 177 (U); Litanie R., Aug 1937, Rombouts 725 (BR, U); 

no locality, Suriname Forestry Bureau 311 (K, U); Biegie 

Stong, 18 Sept 1949, Suriname Forestry Bureau 603 (K, U); 

Coppename R., 17 May 1950, Suriname Forestry Bureau 704 

(U); upper Suriname R., 16 June 1921, Suriname Forestry 

Bureau 5462 (U); Suriname R., 10 July 1908, Tresling 96 (U), 

102 (L), near Goddo, 19 Aug 1908, 33f (RB, U); Gonini R., 

.4ug 1903, Versteeg 157 (U); Tapanahoni R., Aug 1904, Versteeg 855 (U); no locality or date, Voltz 5.71. (U); lower Coppename R., Aug 1901, Went 116 (U); Para, 1852, Wullschlagel 

149-A (BR); Jodensavanna, upper Suriname R., 1852, Wullschlagel I50 (BR), 2069 (W). FREKCH GUIANA: Comte 

R., Belizon, Apr 1961, Aubreville 313 (P); Balate Creek, 23 

Sept 1954, BAFOG 219M (P, U); Criques Ouaqui, Grand & 

Petite, 7-13 Sept 1961, BAFOG 7831 (U); Itany and Marouini, 

29 Sept-2 Oct 1961, BAFOG 7937 (U); Saut des Polissoirs, 

Lunier R., 1897-98, Geay 5.72. (P); Saut Grand Bacon, Mana 

R., Halle 732 (P, US); Rio Oiapoque, across from Colonia 

Agricola do Oiapoque, 4 km north of mouth of Cricu River, 

13 Aug 1960, Irwin et al 47471 (NY, U, US); no locality, 1792, 

Leblond 5.n. (G); no locality or date, Leprieur 5.n. (G, P, US); 

no locality or date, Martin s.n. (BM), Melinon 102 (P); no 

locality, 1821, Perrottet 23 (G); no locality or date, Perrottet 

s.n. (P), Poiteau 5.71. (G, K); no locality, 1863, Rech 45 (P); 

no locality, 20 Oct 1938, Vaillant 5.72. (P). BRAZIL. Para Rio 

hfapuera, 3 Dec 1907, Ducke 9022 (G, RB); Counany, 22 Oct 

1895, Goeldi 1158 (BM, G, RB, US). Terr. do Amapa: 

Oiapoque R., 1 Oct 1949, Black 49-8301 (GH, INPA, NY); 

Oiapoque, Rio Oiapoque, Black et al 54-17621 (IAN); Parque 

Indigena do Tumucumaque, Rio Paru de Oeste, Missa0 

Tiriyo, 27 Feb 1970, Calvalcante 2538 (NY); Rio Oiapoque, 

Aug 1949, Candido s.n. (IAN); Rio Oiapoque, 29 Jan 1950, 

Froes 25698 (IAN) and 15-16 Oct 1950, Froes 26638 (IAS, 

US), 26639 (US), 26649 (I.4N, RB, US); no exact locality or 

date, Luetzelburg 20211 (M, 1%'); Oyapock, Salto Guama, July 

1927, Luetzelburg 20399 (M); Rio Oiapoque near base of 

granite outcrop near Mt. Carupina, 15 Oct 1960, Pires LTVestra 48849 (F, NY, US); Rio Araquari between Monguba 

and Serra do Navio, 25 Sept-4 Oct 1961, Pires et al 51182 

(NY, U, US), 51501 (NY, US); Valle da Selra Tumuc-Humac, 

3 Dec 1928, Sampaio 5760 (R); Paru do Cumina, 7 Dec 1928, 

Sampaio 5818 (R). 

LOCAL NAMES AND UsEs.-Apa (Irwin & Westra 

47471), oeverwallaba (Maguire et a1 54014), ouapa 

(Halle 732), walaba (Lanjouw 1209), wapa rouge 

(Lemee s.n.), watafa (Guppy 198). 
NUMBER 28 25 

3b. Eperua rubiginosa Miquel 

var. grandijlora Pulle 

FIGURE 11 

Eperua rubiginosa Miquel var. grandipora Pulle, Enum. Vasc. 

Pls. Surinam 211, 1906. 

Stipules broadly oval, 20 mm long and 17 mm 

wide, or ovate-triangular, 4 mm long and 3 mm 

wide, acute; leaf rachis (11-)15-22 mm long, the 

leaflets 4- or 5-jugate, the blades more or less inequilateral basally, the lower side somewhat shorter 

and more acute, at least the upper two or three 

pairs; inflorescences 6.5-8.5 dm long, the pedicels 

mostly 2.5-3 mm in diameter; sepals 21-35 mm 

long, 7-15 mm wide. 

TYPE COLLECTION.-G. M. Versteeg 48 (holotype U), “fluv. Gonini” Suriname, August 1903. 

D1sTRIBuTIoN.-Rain forest of interior Suriname 

at low elevations. 

SURINAME: Gonini R., 25 Feb 1918, Gonggrijp 3700 (U); 

no locality, 1841, Hostmann 129 (BM) and 229-A (BM, G, K); 

Jacob Kondre, Saramacca R., 16 June 1944, Maguire 23804 

(A, F, U, US); Coppenam R., Onobissi, 15 March 1916, 

Suriname Forestry Bureau 2577 (NY). 

Hostmann 229 is the number of the type collection of E. rubiginosa Miquel, so the collection cited 

above, showing all the characters of variety grandifiora Pulle, will be cited with an “A’ suffix to distinguish it. It surely represents a mixture of collections somewhere at some time; otherwise one 

must assume that there are several collections with 

flowers exhibiting some sort of abnormality in the 

typical population. Such an assumption has not 

been borne out by dissections of the flowers. 

* leucantha 

FIGURE 1 1.-Geographic distribution of several Eperua species. 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

DrscussIoN.-until new field study brings together more adequate representation of the two 

forms of E. rubiginosa, varieties rubiginosa and 

grandiflora will be maintained separately. They 

differ in flower size and in number of pairs of leaflets. The species differs from all the others with the 

elongate pendent inflorescences in its unique (for 

this genus) multicellular, irregularly branching 

pubescence (Figure la) and from most of its relatives in having nonfalcate leaflets. It shares that 

characteristic with E. glabra, perhaps its nearest 

relative, which besides being almost totally glabrous has eglandular sepals. The two species also 

have all stamens fertile, as well as lateral (rather 

than intrapetiolar) stipules. The verrucose pollen 

of this species appears to be intermediate between 

the punctate-rugulose pollen of the basic stock and 

the punctate form of more advanced taxa. 

4. Eperua falcata Aublet 

FIGURES 5, 13 

Eperua falcata Aublet, P1. Guian. 1:369, t. 142, 1775. 

Punzera falcata (Aublet) Willdenow, Sp. P1. 2(1):540, Mar 

1799. 

Dhorpha falcata (Aublet) J. E. Smith in Rees, Cyclopedia 

11(2), 28 Nov 1808. 

Large tree 8-40 m tall, the trunk 20-80 cm in 

diameter, the bark brownish-gray with rather large 

lenticels, the branchlets minutely puberulous, 

usually glabrescent and often nitid; stipules caducous, 2.5-4.5 mm long, 1.5 mm wide, triangularlanceolate, acute, joined in a bifid, intrapetiolar 

stipular body, glabrous or minutely puberulous 

externally; petioles (23-)30-40 (-50) mm long, 

usually minutely puberulous, the rachis (32-)50-75 

(-85) mm long, usually minutely puberulous; leaflets 3- or 4-jugate, (seedlings bijugate), the petiolules 

(3-)4-5 (-6) mm long, canaliculate on the upper 

side, minutely puberulous, sometimes glabrous or 

glabrescent, the blades minutely pellucid-punctate, 

plane, chartaceous to thinly coriaceous, glabrous, 

falcate-elliptic, (7-)9-12 (-15) cm long, 3-7.5 cm 

wide, the base usually obtuse or rounded-obtuse, 

infrequently acute, the apex acuminate to longacuminate and usually mucronulate, the costa and 

venules prominulous; inflorescence a terminal, 

pendent raceme of racemes (6.5-)lo-I4 (-25) dm 

long, the axis usually glabrous except minutely 

strigulose in the floriferous portion, the hairlets 

yellowish-brown, the lateral racemes (27-)30-50 

(-100) mm long, the bracts caducous, 2-3.5 mm 

long and wide, broadly ovate to semicircular, strig 

ulose externally, glabrous on inner surfaces, the 

bracteoles caducous, 3-4 mm long, 3-5 mm wide, 

broadly ovate, pubescent as bracts, arising near the 

base of the hypanthium; pedicels 9-18 mm long, 

minutely strigulose, the hypanthium 4-5 mm long, 

5 (-8) mm wide, minutely strigulose externally; 

sepals deep red to red-brown, pink within, 16-20 

mm long, 6-14 mm wide, oblong to oval-oblong, 

obtuse, minutely strigulose externally, glabrous 

within, the two outer sepals broader, cucullate; 

petal pink to scarlet, 13-18 mm long, 19-25 mm 

wide, oblate-flabelliform, basally truncate, glabrous, 

the 4 petalodia semicircular, 1 mm in diameter, 

glabrous; filaments of 4 or 5 fertile stamens pink, 

united with 4 or 5 sterile (the anthers smaller) 

ones in a villose to villose-sericeous, equilateral 

tube ca. 4 mm long, the filament of the one free 

stamen villose in the basal half, 20-30 mm long, 

filaments of the larger stamens 32-45 mm long, 

glabrous above stamen tube, smaller ones 20-25 

mm long, villose, the anthers ca. 8 mm long and 

3.5 mm wide, oval-oblong, glabrous; stigma capitate, the style glabrous, 40-42 mm long, the ovary 

oblong to oblong-obovate, 6-8 mm long, 3 mm 

wide, velutinous, the gynophore 2-5 mm long, puberulous to glabrous; fruits flat, somewhat arcuateoblong, 24-36 cm long, 6.5-9.5 cm wide, minutely 

strigulose but eventually glabrous, the seeds flat, 

oval, 33 mm long, 22 mm wide, the margins incrassate, the seed coat thin, red-brown, minutely 

tessellate. 

TYPE COLLECTION.--F. Aublet s.n. (lectotype 

BM) “Cayenne 1775.” The lectotype sheet has 

created some confusion and could easily affect the 

nomenclature of the species. The specimen consists of an inflorescence representing Aublet’s species, unattached to vegetative material of another 

species altogether, probably E. rubiginosa. There 

are BM sheets collected by Martin in “Cayenne” 

that bear mixtures of E. rubiginosa and E. falcata, 

as well as several which have only material of the 

first of this pair. Aublet’s plate and description 

surely represent what has been called E. falcata 

by most workers in recent generations. Thus, the 

lectotype must be restricted to the inflorescence on 

the BM sheet. 

DIsTRIBuTI0N.-Frequent to dominant in riv- 
NUMBER 28 27 

erine or low elevation, dry evergreen or swampwoodland rain forest on a wide range of soils 

(often pure white sand) at elevations up to 700 m 

in southeastern Venezuela, central Guyana and 

Suriname, coastal French Guiana, and adjacent 

northeastern Brazil; planted in the Trinidad Botanic Gardens, Cameroon (West Africa), and in 

West Java in the Bogor Botanic Gardens. (The collection from the Victoria Botanical Garden, Cameroon, bears a binomial-“Eperua preussii”-which 

I have not found in any of the standard floras, Index Kewensis, etc. It apparently has no validity). 

Flowering from July to January; fruiting August 

to April. 

VENEZUELA. Bolivar: Between El Dorado and km 88, 9 

Sept 1966, Blanco 603 (US); 90 km from El Dorado toward 

Santa Elena, 28 Dec 1956, Foldats 2645 (US); between kms 

42 and 65, road south of El Dorado, 230 m, 26 July 1960, 

Steyermark 86692 (NY, US). GUYANA: Morebo, Barima R., 

11 Dec 1908, Anderson 122 (K); Botanic Gardens, Georgetown, Oct 1904, Bartktt 8150 (NY); upper Rupununi R., 

near Dadanawa, 28 May 1922, de la Cruz 1373 (F, GH, NY, 

US); Essequibo-Mazaruni Divide, 2 Sept 1932, Davis 319 (FD 

2312) (K); Lima, west bank of Essequibo R., 4 May 1960, 

Harris TP-404 (K); Corantijne R., Oct 1870, Im Thurn s.n. 

(K); Corantijne R., Nov 1879, Jenman 234 (K) upper Demerara R., Sept 1887, Jenmalt 4095 (K), 4101 (K), Nov 1888, 4941 

(K); near Bartica, Nov 1888, Jenman 5015 (BM); Mazaruni 

R., Lung 276 (NY); Comaka, Demerara R., May 1923, Persaud 243 (F); Mazaruni R., Lung L- Persaud 272 (F); Essequibo 

R., Moraballi Creek near Moraballi-Wineperu Divide, 28 Aug 

1929, Sandwith 137 (K, NY, RB, U), 23 Sept 1929, 329 (K, 

RB, U, US); Trail Casobrikeva to Pipilipai, Kukui R., 6 

Sept 1960, Tillett G Tillett 45304 (F, NY, RB, US); near falls 

of Kako R., 23 Sept 1960, Tillett G Tillett 45413 (NY); 

Kamarang R., west bank above Utschi R., 550 m elev., 25 

Oct 1960, Tillett LY Tillett 45791 (NY, RB) and 45794 (GH, 

NY, US). SURINAME: Upper Coppename R., Sept 1901, 

Boon 1199 (U); Vosberg, Caronie R., 1947, Burger 8 (U); 

Jodcnsavanne-Mapane Creek area, 9 Dec 1963, Elburg 9822 

(U); Sarakreek, near Sinnannamary, 13 h’ov 1950, J. Lr P. A. 

Florschutz 227 (U); “Boven Para,” Sept, Focke 845 (U); no 

locality or date, Hostmann s.n. (M); Gran Rio R., 6 Sept 

1910, Hulk 227 (U); 1-5 km north of Lucie R., 3 km west 

of Oost R., alt 225 m, 8 Sept 1963, Irwin et al 55448 (GH, 

NY, U, W); Lucie R., 2-5 km below the confluence with the 

Oost R., 8 Sept 1963, Irwin et al 55467 (F, NY, RB, U, US); 

Zuid R., 2 km above junction with Lucie R., 15 Sept 1963, 

Irwin et al 55716 (GH, NY, U, US); Fallawatra 3 Nov 

1971, Jimenez-Saa 2542 (K, U), 11 Nov 1971, 1597 (K, U), 

16 Nov 1971, 1622 (K) and 1625 (U), 17 h’ov 1971, 1627 (K); 

Jodensavanne-Mapane Creek area, Suriname R., 19 Sept 

1953, Lindeman 4649 (F, U); base of Tafelberg, 13-15 

Sept 1944, Maguire 24720 (A, F, NY, U, US) and 24778 

(A, F, NY, U, US); Frederik Top, 2 km SE of Juliana Top, 

700 m elev., 29 July 1963, Maguire et al 54401 (F, G, GH, M, 

NY, U); Tanjimama R., 17 Nov 1954, Mennegu 441 (U); no 

locality, 1850, Miquel s.n. (K); Mapane region, 7 Nov 1967, 

Sabajo Q Roberts 11192 (U); Juliana Top, 550 m, 29 July 

1964, J. P. Schulz 10237 (U); no locality or date, Schweinitz 

s.n. (NY); Sectie “0,” 28 Oct 1915, Suriname Forestry Bureau 

1171 (U), 7 Feb 1916, I609 (U), 17 March 1917, 2750 (U), 26 

July 1917, 3100 (U), 1 Nov 1917, 3385 (BR, U), 6 Nov 1917, 

3401 (U, US), 5 Kov 1917, 3408 (NY, U), 26 Jan 1918, 3618 

(K, U), 28 Jan 1918,3625 (U); 12 Aug 1918, 3970 (BR, U, US), 

17 Aug 1918, 3972 (K, U), 5 Dec 1918, 4192 (U), 9 Dec 1918, 

4200 (U), 16 Sept 1919,4416 (NY, U), 22 Dec 1919, 4528 (U), 3 

Oct 1920, 4791 (U), 7 Feb 1921, 5042 (U), 3 Nov 1921, 5408 

(U), 3 h’ov 1921,5420 (U), 24 Nov 1922,5999 (U); Brownsberg, 

10 Apr 1916, Suriname Forestry Bureau 1725 (U), 10 Oct 1917, 

3210 (U), 22 Nov 1917, 3434 (U), 27 Sept 1919, 4420 (K, NY, 

U), 15 March 1922, 4536 (U), 10 Sept 1921, 5364 (U, US), 26 

Oct 1921, 5547 (U), 18 July, 6906 (BR, U); Zanderij I, 8 Nov 

1918, Suriname Forestry Bureau 4061 (U), 28 May 1919, 4324 

(U), 14 Oct 1920, 4838 (U), 27 Jan 1921, 5060 (NY, U), 28 

Apr 1922, 5804 (U); Watramiri, 11 Oct 1921, Suriname Forestry Bureau 5398 (U), 19 Dec 1921, 5549 (U); Blakhawatra, 

28 Oct 1921, Suriname Forestry Bureau 5524 (U); Litanie R., 

near Knopaiamoi, Dec 1903, Versteeg 393 (U); Zanderij I, 

Oct. 1944, Woodherbarium Suriname 22 (A, K, NY, U), 23 

(A, K, NY, U); Sectie “0,” Sept 1942, Woodherbarium Suriname 66 (A, K, NY, U); Paramaribo, Wullschliigel 149 (NY, 

W); Para, Wullschliagel s.n. (BR). FRENCH GUIANA: St. 

Laurent, Maroni R., 14-15 April 1961, Aubreville 236 (P, U) 

and 260 (US); route de St. Laurent-Cayenne, km 11, 7 Aug 

1953, BAFOG 52M (P, U); Crique Serpent, 2 Feb 1954, 

BAFOG 143M (P, U): Cayenne, Nov-Dec 1956, BAFOG 1274 

(U); placeau no. 2, carreau no. 57, route de Mana, 22 Nov 

1955, BAFOG 7062 (NY, P, U), 23 Nov 1955, 7064 (NY, P, U); 

St. Laurent, Feb 1956, BAFOG 7254 (U) and 7298 (U); Charvein, Trace de L’Acarouany, 10 Nov 1913, Benoist 193 (P); 

Charvein, 16 Jan 1914, Benoist 596 (P); upper Maroni, 1877, 

Crevaux s.n. (P); Mana R., Crique Tamanoir, 18 Aug 1962, 

Hulk 579 (P); 0.5 km south of mouth of Rio Matura 21 

Sept 1960, Irwin et al 48408 (GH, NY, U, US): Camopi R., 

near junction with Oiapoque R., 2 Oct 1960, Irwin et al 

48613 (GH, NY, U, US); Maroni, Charvein, Mar 1901, Lemee 

s.n. (P); Cayenne, Martin s.n. (BR, K); no locality or date, 

Melinon s.n. (F, GH, K, NY, P, R, US); Balate, St. Jean du 

Maroni, 23 Sept 1970, I. Petrov 239 (P); no locality or date, 

Richard s.n. (G, W); Karouany, 1896, Sagot 183 (BM, G, K, 

NY, P, W); no locality or date, Vaillant s.n. (US); Maroni, 

Wachenheim 130 (P): Godebert, Wachenheim 405 (US). BRAZIL. Amapa: Rio Oiapoque, 19 Oct 1950, Froes 26687 (IAN, 

NY, P, RB, US); edge of Rio Ingarari near junction with Rio 

Oiapoque, 15 Sept 1960, Irwin et al 48277 (F, NY, US); Rio 

Oiapoque near Mt. Carupina, 16 Oct 1960, Pires et al 48876 

(NY, US). 

LOCAL NAMES AND UsEs.-Apa (Irwin et a1 

48277), apa roxo (Pires et a1 48876), baboen walaba (Woodherbarium Suriname 66), bioudou 

(FGFD 7062), biouolou wata (BAFOG 143M), 

bootlace tree (Broadway s.n., Trinidad), bylhout 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(Focke 845), caraota (Blanco 603), itoeli walaba 

(Woodherb. Sur. 66), mahomillo negro (Steyermark 86692), rosa de montafia (Foldats 2645), 

sand wallaba (Harris TP-404), tamoeno pale-o 

(Woodherb. Sur. 23), soft wallaba (Sandwith 329), 

wapa gras and wapa huileux (BAFOG 1274), wallaba (Broadway s.n.), walaba koeleroe (Woodherb 

Sur. 23), white wallaba (Davis 319), zwarte walaba 

(Woodherb. Sur. 22); waraba (Tillett & Tillett 

45794). Used for making shingles. 

DIscussIoN.-The most frequent, widely distributed species of the genus, E. falcata, is easily distinguished from the other two species with falcate 

leaflets and elongate, pendent inflorescences (E. 

venosa and E. leucantha). Its brownish-goldcolored, minutely strigulose inflorescences and sepals, its villose staminal tube, and its capitate stigma (rather than bilobed-capitate) serve to separate 

it in field or herbarium. From E. rubiginosa, with 

which it is sometimes confused, it differs, among 

other things, in the eglandular outer sepals; E. 

rubiginosa has an apical-dorsal gland on each of 

the two outer sepals. Palynologically, E. falcata is 

unique by virtue of its heterobrochate-reticulate 

ektexine. Although two other species, E. purpurea 

and E. oleifera, have pollen-ektexines that are classified as reticulate, comparison of the scanning 

electron micrographs (Figure 9) reveals two distinctive ektexine patterns. Taking all data into consideration, E. falcata is nearer the other pendentinflorescence species, especially E. ru biginosa, than 

to the E. purpurea-E. oleifera alliance. 

5. Eperua venosa Cowan 

FIGURES 5, 11 

EFerua venom Cowan, Mem. N. Y. B. G. 10:148, 1 July 1958. 

Large tree to 30 m tall and 60 cm in diameter, 

the branchlets glabrous; stipules persistent, rigid, 

connate laterally in a compound, bifid, intrapetiolar body, 4-5 mm long, ca. 2 mm wide, lanceolatetriangular, acute, glabrous externally but sericeous on the inner surface, ciliolate; leaves glabrous, 

lucid, the petioles (22-)32-45 mm long, the rachis 

30-55 mm long, terete; leaflets bijugate, the petiolules (6-)9-13 mm long, the blades pellucidpunctate, plane, coriaceous, (1 1.5-) 14-22 cm long, 

(5-)8-9 cm wide, elliptic to broadly elliptic, the 

apex abruptly or gradually acute to acuminate, 

sometimes mucronate, the base of the lower pair 

rounded-obtuse, the upper pair inequilateral basally, obtuse on the lower side but attenuate-acute 

on the upper side, the venation prominently salient on both blade surfaces; inflorescence a pendent, terminal raceme of racemes (85-)145-188 cm 

long, the axes minutely puberulous, the lateral 

racemes 2-4.5 cm long, the bracts early caducous, 

about 3 mm long and 3.5 mm wide, broadly ovate, 

ciliolate, strigulose externally, glabrous on the 

inner surface, the bracteoles caducous, arising in 

the upper part of the pedicel, ca. 4-4.5 mm long 

and 4 mm wide, strongly concave, broadly ovate, 

pubescent as bracts; pedicels 8-15 mm long, graysericeous, the hypanthium 4-7 mm long, 6 mm in 

diameter, minutely sericeous externally, the nectaries projecting just above the hypanthium rim; 

sepals green with purple tint, the two outer ones 

broader and cucullate, 20-30 mm long, 9-13(-17) 

mm wide, oblong to elliptic-oblong, obtuse, minutely sericeous outside, glabrous within; petal 

deep rose-colored, 25-34 mm long, 48-60 mm wide, 

broadly oblate-flabelliform, strigulose externally at 

the base, glabrous within, the four petalodia ca. 

0.8 mm long and wide, semicircular, the margin 

crenulate; 10 stamens fertile, filaments 40-50 mm 

long, the staminal tube 8-10 mm long, somewhat 

arcuate, densely sericeous on the inner and outer 

surfaces, the base of the free portion of the sterile 

filaments also sericeous, the anthers 6.5-9 mm long, 

3-3.5 mm wide, glabrous; stigma capitate, sometimes obviously bilobed, the style 45-60 mm long, 

glabrous except at the extreme base, the ovary 

oblanceolate, 8-10 mm long, 3.5-4.5 mm wide, 

densely pilose-sericeous, the gynophore pilosesericeous, 3.5-6 mm long; fruit not seen. 

TYPE COLLECTION.-H. R. Kunhardt, Jr., 20 

(holotype NY), Angel Falls, Bolivar, Venezuela. 

DIsTRIBuTIoiv.-Frequent in lowland rain forest 

in southeastern Venezuela. 

VENEZUELA. Bolivar: El Foco, Rio Uiri-yuk, 30 August 

1962, Maguire et al 53514 (NY, US); 'Triana Savanna, Cerro 

Pioh, 5 Sept 1962, Maguire et al 53597 (BR, NY, U, US). 

DISCUSSION.-A cluster of three species-E. venosa, E. leucantha, and E. falcata-may have been 

derived from the E. rubiginosa-E. glabra group. 

All three have falcate leaflets and bifid, nonfoliaceous, intrapetiolar stipules. The ten fertile stamens of E. venosa link it to E. rubiginosa-E. glabra, 
NUMBER 28 29 

but a closer relationship is evident with E. leucantha, from which E. venosa differs in petal color 

(deep rose vs. white) and pubescence, as well as 

in number, form, and venation of the leaflets. The 

fine ashy puberulence of the inflorescence and sepals in E. zlenosa and E. leucantha contrasts with 

E. falcata, as well as with that of E. rubiginosa. The 

petal of E. venosa is somewhat attenuate and externally puberulous basally, rather than truncate 

and glabrous as in all the nearly related species. 

The pollens of E. venosa and E. leucantha are very 

similar in being the only two that are 3-porate as 

well as punctate-rugulose. 

6. Eperua leucantha Bentham 

FICURFS 6, 11 

Eperua leucantha Bentham, F1. Bras. 15(2):225, 1870. 

Small to large tree (8-)12-25 m tall, (10-)50-60 

cm in diameter, the bark gray, smooth, the branchlets micropuberulous at first, glabrescent; stipules 

deciduous, lanceolate, sometimes somewhat arcuate, joined in an intrapetiolar body 3-5 mm long, 

1.5-2 mm wide, glabrous except marginally ciliolate; petioles (17-)25-40 (-55) mm long, glabrous 

or infrequently micropuberulous and glabrescent, 

the rachis (27-)50-95 (-123) mm long, glabrous 

or pubescent as petioles; leaflets mostly 3-jugate, 

sometimes 2- or infrequently 4-jugate, the petiolules (5-)7-8 (-10) mm long, usually glabrous but 

sometimes micropuberulous, sulcate on upper surface, the blades pellucid-punctate, (8-)10-16 (-20) 

cm long, 3.5-7 (-8.5) cm wide, chartaceous to 

thinly coriaceous, plane, falcate-elliptic, the base 

mostly rounded-obtuse or obtuse, sometimes the 

upper pairs acute basally, infrequently the lower 

side of the leaflet obtuse and the upper side 

attenuate-acute, the apex acuminate to longacuminate, the tip often obtuse and usually mucronate, glabrous or rarely micropuberulous at the 

base of the costa on the upper surface, the venation prominulous to subobscure; inflorescence a 

terminal, pendent raceme of racemes (37-)60-90 

(-1 16) cm long, the main axis micropuberulous, 

the axis of the lateral racemes (6-)7-8 (-10) mm 

long, microstrigulose, the bracts caducous, the bracteoles arising about midway on the pedicels, caducous; pedicels (5-)lo-14 (-22) mm long, micropuberulous to microstrigulose, the hairlets grayish, arcuate, the hypanthium (3-)4-5.5 mm long, 

5-7 mm in diameter, microstrigulose externally, 

the nectaries slightly exserted, inconspicuous; sepals pellucid-punctate, concave, fleshy, white, oval 

to oblong, obtuse, two outer ones broader and cucullate, (15-)18-20 (-29) mm long, 7-10 (-12) mm 

wide, micropuberulous or microstrigulose externally, inner surfaces glabrous; petal white, 20-33 

mm long, 36-55 mm wide, broadly flabelliform, 

basally truncate, sessile, glabrous, four petalodia 

0.8-1.5 (-5) mm long, 0.8-1.5 mm wide, mostly 

semicircular to oblate or oval; all 10 stamens ap 

parently fertile, filaments joined at the base in a 

subequilateral tube 3.5 to 5 mm long, the tube and 

the base of shorter filaments sericeous externally 

and on the inner surfaces, the longer stamens 38- 

56 mm long, the shorter ones 30-43 mm long, the 

anthers 5-8 mm long, 2-3 mm wide, oval to oblong, 

the thecae microtessellate; stigma capitate, sometimes somewhat bilobed, the style 33-53 mm long, 

glabrous, the ovary sericeous-velutinous, 6-10 mm 

long, 3-4 mm wide, oblong-oblanceolate or oblongobovate, the gynophore sericeous, 2-5 mm long; 

fruit 25-31 cm long, 6.5-9.5 cm wide, minutely 

strigulose, seeds oval, flat, 35 mm long and 21 mm 

wide. 

TYPE COLLECTION.--R. Spruce 2021 (holotype 

K, isotypes BM, G, GH, M, P, W), "Rio Negro, 

moist forest at Uanauaca, Jan 52," Amazonas, 

Brazil. 

On the holotype label, Spruce records the following: 

This tree, called Jauacanza in Lingoa Geral grows throughout 

the Upper Rio Negro, Uaupes and Casiquiare, where along 

with the Jeharli [Eperua purpurea] it is characteristic of the 

caatingas, and is perhaps the more frequent of the two. It 

flowers nearly throughout the dry season, but is in greatest 

perfection in the months of September & October. No use 

seems to be made of any part of it, but the flowers afford a 

great deal of honey to bees, as do also those of two species 

of Humirium growing in the driest parts of the caatingas 8c 

flowering at the same time. When the Indians see these trees 

flowering copiously they predict an abundance of wild 

honey. 

DISTRIBUTION.-FreqUent in uppermost Rio 

Negro Basin in Brazil, west to the Rio Vaupes 

drainage of southeastern Colombia, east and 

north to the Rio Guainia-Rio Casiquiare in southwestern Venezuela on sandy soils sometimes in 

savanna-border forest, at 120 to 250 m elevation; 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

flowering October to January; fruiting February 

to April. 

COLOMBIA. Vaupes: San Felipe, Rio Negro, 13-25 Nov 

1952, Humbert 27386 bis (US); Rio Kuduyari, 16 Oct 1952, 

Schultes b Cabrera 17872 (BM, GH, NY, U, US). VENEZUELA. Amazonas: Yavita-Pimichin Trail, near Pimichin, 21 

Nov 1953, Maguire et al 36307 (NY, US); Rio Yatua near 

Cano Tauavaca, 4 Dec 1953, Maguire et al 36508 (F, NY); 

Rio Yatua near Cerro de la Neblina, 3 Jan 1958, Maguire 

et al 42610 (F, GH, NY, RB, U, US); San Carlos de Rio 

Negro, 17-18 Apr 1970, Steyermark A+ Bunting 102733 (US); 

Yavita, 23 Jan 1942, LI. Williams 13939 (A, G, RB, US); 

Maroa, Rio Guainia, 17 Feb 1942, LZ. Williams 14390 (F, G, 

US); Pimichin, Rio Guainia, 3 Apr 1942, Ll. Williams 14940 

(G), 1 Apr 1942, 14942 (F, US); Capihuara, alto Casiquiare, 

5 June 1942, Ll. Williams 15801 (F, G, US). BRAZIL. Amazonas: Sao Gabriel, Rio Negro, 29 Oct 1932, Ducke 164 (A, 

F. NY); Rio Negro above mouth of Rio Curicuriary, 19 Nov 

1929, Ducke 23289 (G, RB, US); Sao Gabriel, upper Rio 

Negro, 28 Oct 1932, Ducke 23732 (P, RB, U); Porto Curucuhy, 

Rio Negro, 6 Oct 1945, Froes 21101 (F, IAN, NY, US); Sao 

Felipe, Rio Negro, 1 Oct 1952, Froes 28826 (IAN); Rio 

Cauaburi, 7 Nov 1965, Maguire et al 60130 (F, NY, US); 

Marabitanas, 20 Apr 1947, Pires 481 (IAN); Rio Negro-Rio 

Vaupes, 1 May 1947, Pires 531 (NY, US); Rio Vaupes, 

Taraqua, Pires 978 (IAN); road from Camanaus to ‘(iaupes, 

30 Oft 1971, Prance et al 15684 (GH, M, NY, U, US); Rio 

Negro, Ilha das Flores, 8 March 1959, Rodrigues &+ Pires 178 

(IAN, US); upper Rio Negro, Ilha das Flores, 17 Feb 1959, 

Rodrigues 905 (U, US). 

LOCAL NAMES AND UsEs.-Acana, IuacanHa(Rodrigues 905), jauacana (Spruce 2021), yaguana, yahuana (Ll. Williams 14390), yauacano (Ducke 23732). 

The wood is very heavy and is used locally in 

l‘enezuela for house posts and for bridges. 

Some of the stamens (two to six) in flowers of 

E. leucantha do not bear fertile anthers, a characteristic shared with E. falcata. The white petal of 

E. leucantha contrasts sharply with all the other 

species with the elongate, pendent inflorescences, 

although less so with E. glabra which has a creamcolored petal. The nearest relative is unquestionably E. venosa which has a deep rose-colored petal 

with the base attenuate and puberulous externally, 

and ten fertile stamens. Their pollens are virtually 

identical (cf. discussion following E. venosa). 

7. Eperua duckeana Cowan, new species 

FKUW 6, 11 

Arbor parva ad mediocris 7-25 m alta, trunco 

20-30 diametro, ramulis glabris et nitidis; stipulae 

plerumque deciduae foliaceae glabrae, 9-1 5 mm 

longae, 9-10 mm latae, inaequilateraliter ovales 

ad ovatae; petioli 10-35 mm longi, glabri, rhachibus (8-)I 1-20 cm longis, glabris; foliola plerumque 4-jugata, petiolulis 4-8 mm longis, plerumque 

glabris, laminis aliquando plus minusve pellucidopunctatis, coriaceis, planis, 6.5-10 (-12) cm longis, 

3-6 cm latis, plerumque uno vel duo jugis basalibus ovatis ad lanceolatis et rotundato-obtusis, 

reliquis jugis ellipticis et basi subobtusis ad acutis, 

acuminatis ad longo-acuminatis mucronulatisque, 

glabris aut raro micropuberulis ad basem supra, plus minusve nitidis supra, venis venulisque 

prominulis; inflorescentiae plerumque terminales 

composito-racemosae, aliquando solum racemosae, 

3-8 cm longae, axe tomentuloso, pilis arcuatis bifidis, bracteis bracteolisque caducis; pedicelli 8- 

20 mm longi, tomentulosi, hypanthio 3-4 mm 

longi, tomentulosi, hypanthio 3-4 mm longo, 4-6 

mm diametro, tomentuloso, nectariis super labrum 

hypanthii procurrentibus; sepala diluto-viridia, 

tomentulosa extus, intus glabra, oblonga ad ovalia 

ad elliptica, 12-18 mm longa, 4-9 mm lata, uno 

dorsuali latiore et concavo, plerumque sepalis externis duobus glandulo versum apicem ornatis; 

petalum album, glabrum aut raro leviter puberulum basaliter, 14-25 mm longum, 22-35 mm latum, 

oblatum, truncatum ad basem, petalodiis 1-2 mm 

longis; omnia stamina fertilia, filamentis 26-62 

mm longis, 9 tubo basaliter conjugentibus, tub0 

plus minusve zygomorpho, 3.5-9 mm longo, tomentuloso et glabro virgato, antheris oblongis ca. 3.8 

mm longis et 1.5 mm latis, glabris; stigma capitellatum, stylo 26 mm longo, glabro, ovario oblanceolato vel oblongo-oblanceolato, 7-8 mm longo, 3- 

3.5 mm lato, dense tomentuloso, gynophoro 6-10 

mm longo, tomentuloso; fructus tomentulosus, 16 

cm longus et 6 cm latus (submaturus), oblanceolatus, planus, carpophoro ca. 20 mm longo, tomentuloso. 

TYPE COLLECTION.--A. Dzicke 732 (holotype 

US sheet 1875576, isotypes F, NY, R) Rio UrubG 

below Iracema Falls, Amazonas, Brazil, 7 July 1941. 

DIsTRIBuTI0N.-Primary forest on sandy or clay 

soil, especially along rivers, but above the annual 

flood levels, in the vicinity of Manaus. Flowering 

June to September; fruiting May to December. 

BRAZIL. Amazonas, vicinity of Manaus: Reserva Florestal 

Ducke, 20 Sept 1968, Aluisio 172 (INPA, US); edge of Riacho 
NUMBER 28 31 

Grande, 15 Nov 1956, Coelho 4308 (US), 15 Nov 1956, 

Dionisio (INPA no.) 4308 (IAN); upper Mindu R., 19 Nov 

1935, Ducke 244 (NY); Estrada do Aleixo, 25 Aug 1931, Ducke 

23734 (G, P, RB, U, US); Rio Taruma, 9 Aug 1949, Froes 

24949 (IAN); Rio Urubu, 20 Oct 1949, Froes 25330 (IAN); 

Estrada BR-17, km 3, 13 June 1956, Melo 2 Coelho 3919 (US); 

Kms 60-70 Manaus-Itacoatiara, 21 Oct 1963, Oliveiru 2737 

(IAN); Estrada AM-1, km 70, 13 May 1969, Osmarino 5.72. 

(INPA, US); Manaus to Caracarai Road, km 26, 10 Nov 1966, 

Prance et al3072 (F, GH, SY, R, U, US); Rio Urubu between 

Iracema Falls and Natal, Caracarai Road, 5 June 1968, Prance 

et a1 4977 (F, G, GH, M, NY, R, U, US); Estrada ManausItacoatiara, km 202, 19 Dec 1966, Prance et al 3720 (NY), km 

69, 8 Sept 1960, Rodrigues 6 Chagas 1725 (US), 5 Oct 1960, 

Rodrigues et a1 1807 (US), km 65, Reserva Walter Egler, 4 

Feb 1970, Rodrigues 8686 (INPA, US), km 55, 18 March 1961, 

Rodrigues 6 Lima 2233 (INPA, US); Reserva Florestal 

Ducke, 1 Dec 1964, Rodrigues & Osmarino 6769 (INPA, US); 

Igarape do Riacho Crande, 14 Aug 1957, TYilliarn 501 (IAN). 

DIscussIoN.-This species shares many character 

combinations with E. schomburgkiana, but there 

are substantial phenotypic differences. Most conspicuous of these is the pubescent ovary and smaller flowers, as well as the geographic distribution. 

It is named to honor Adolpho Ducke, prodigous 

pioneer in the collection and study of Brazilian 

tree legumes. 

8. Eperua glabrijlora (Ducke) Cowan, 

new combination 

FICURFS 7, 12 

Eperiia bijuga Martius ex Bentham var. glabriflora Ducke, 

Trop. Woods 62:27, 1940. 

Tree 5-35 m tall, the trunk 15-30 cm in diameter, the bark thin, slightly scaly, the branchlets 

glabrous, nitid; stipules 2-4 mm long, 2-3 mm 

wide, triangular to triangular-lanceolate, acute, 

glabrous; leaves glabrous, the petioles (1 2-) 15-25 

(-30) mm long, the rachis (30-)50-75(-85) mm 

long; leaflets bijugate, the petiolules 6-12 mm 

long, the blades somewhat pellucid-punctate, coriaceous, sometimes rigidly so, the margins revolute, 

7-20 cm long, 5-10 cm wide, the lower pair often 

smaller and ovate to elliptic, the upper pair elliptic, the base of all leaflets rounded-obtuse or those 

of the upper ones acute, the apex acute to acuminate, mucronate, nitid on both surfaces, the venation prominulous with the costa depressed on the 

lower blade surface; inflorescence an axillary or 

terminal raceme of racemes (2.5-)7-9 (-10.5) cm 

long, glabrous, the bracts and bracteoles caduceus, 

glabrous, the bracts 5.5 mm long and 6 mm wide, 

broadly ovate, the apex gland-tipped dorsally, the 

bracteoles 7.5-10 mm long, 8 mm wide, oval-ovate, 

acute, gland tipped dorsally near the apex; flowers 

glabrous except for the androecium, the pedicels 

3-10 mm long, glabrous, the hypanthium 2-3 mm 

long, 3-5 mm in diameter, glabrous, the nectaries 

obvious; sepals purple, pellucid-punctate, 13-22 

mm long, 3-10 mm wide, polymorphic, oval, elliptic, oblong, falcate or not, the dorsal one larger, 

concave, the outer two with a gland near the apex 

on the outer surface; petal red to purple, 4-5 cm 

long, 4.5-6.5 cm wide, broadly obcuneate, the base 

attenuate, the petalodia 0.6-1.5 mm long; stamens 

all fertile, 20-45 mm long, 9 united basally in an 

inequilateral tube 7 mm long on the shorter side 

and 10-12 mm long on the other, the tube puberulous externally and on the inner surfaces, the anthers oval 2.5-3 mm long and 1.5-1.7 mm wide; 

gynoecium glabrous, the stigma obtuse, the style 

24-33 mm long, the ovary obcuneate, 4-7 mm long, 

3-3.5 mm wide, the gynophore 5-9 mm long; 

fruits flat, woody, broadly oblong, 12.5-15 cm long, 

6-10 cm wide, the dorsal margins incrassate, the 

carpophore 15-25 mm long, 

TYPE COLLECTION.--A. Ducke 158 (lectotype US, 

sheet 1693166, isolectotypes A, F, NY, R), Estrada 

do Aleixo, 3 April 1936, Amazonas, Brazil. ‘This is 

one of three syntypes cited in the protologue. A 

second collection labeled “158 (11 a Col.)” at IAN 

was taken at the type locality but on 25 June 1944. 

DIsTRlBuTI0N.-Common to dominant in upland, nonflooded rain forest on marshy soils of 

mixed sand and humus in northwestern Brazil, 

especially near Manaus. Flowering from February 

to June; fruiting July to December. 

BRAZIL. Amazonas, Manaus: Estrada Manaus-Itacoatiara 

km 27, 7 Apr 1967, Byron h Elks 67 (INPA, US), June 1932, 

Ducke I4a (A, F), Rio Taruma, 7 Aug 1949, Froes 24926 

(IAN), 21 May 1903, Goeldi 3862 (BM, G, RB, US); 12 Feb 

1958, INPA 6065 (US); Parque 10, 28 March 1958, INPA 

6280 (IAN, US); along road to Aleixo, Aug-Sept 1936, Krukoff 

8006 (BM, BR, G, GH, P, NY, U); km 202, near Rio Urubu, 

Itacoatiara Highway, 19 Dec 1966, Prance et al 3709 (F, G, 

GH, NY, R, U, US); beside Igarape Taruma, 14 March 1967, 

Prance et a1 4637 (F, GH, NY, R, U, US); Reserva Florestal 

Ducke, picada PE, P.25, 26 Aug 1957, Rodrigues 562 (US); 

cachoeira alta do Taruma, 27 July 1961, Rodrigues 6 Chagas 

3061 (US): Reserva Florestal Ducke, near 422, 29 Feb 1964, 

Rodrigues ~9 Osmarino 5723 (US), near Q20, 5982 (us); 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Manaus, Res. F1. Ducke, Quadrat 21, 7 May 1965, Rodrigues 

&+ Osmarino 6932 (INPA, US); 6 Aug 1965, Rodrigues 6 

Osmarino 7005 (INPA, US). Para: Faro, 27 Jan 1927, Ducke 

20313 (RB, U). 

LOCAL NAMES AND USES.-Muirapiranga (Ducke 

158), muirapiranga da folha grande (Rodrigues 

& Osmarino 5982). Frequently used for firewood, 

particularly because the wood is resinous. 

DIscussIoN.-Originally described as a variety of 

E. bijuga by Ducke, E. glabriflora amply deserves 

recognition as a distinct species by virtue of its 

regular hypanthium, glabrous gland-tipped outer 

sepals, longer bracteoles, and most obviously by the 

revolute leaflets. Both species have all ten stamens 

fertile, which character alone separates them from 

all the others with short, erect inflorescences except 

E. schomburgkiana, which has a white petal and 

more pairs of leaflets. There seems little reason 

to doubt the close relationship of E. glabriflora 

to E. bijuga, both having close affinity with E. 

jenmanii. 

9. Eperua bijuga Martius ex Bentham 

FICURFS 7, 12 

Eperua bijuga Martius ex Bentham, F1. Bras. 15(2):226, 1870. 

Eperua bijuga Martius ex Bentham forma typica Ducke, 

Trop. Woods 62:26, 1940. 

Small tree 7-12 m tall, the branchlets glabrous, 

shiny; stipules foliaceous, free from each other, 

falcate-elliptic to falcate-oval, 7-12 mm long and 

3-10 mm wide, persistent, rigid, glabrous; leaves 

glabrous, the petioles 8-17 (-23) mm long, the 

rachis 25-65 mm long; leaflets bijugate, the petiolules 6-1 3 mm long, the blades pellucid-punctate, 

plane, coriaceous, 9.5-20.5 cm long, 3.5-8.5 cm 

wide, elliptic to ovate or lanceolate, the lower 

pair sometimes shorter, ovate-elliptic and basally 

rounded-obtuse, the upper pair elliptic and basally 

acute, the apex acuminate or acute, sometimes 

bluntly so, the venation subobscure; inflorescence 

an axillary or terminal raceme or raceme of racemes 1.5-4 cm long, the axes puberulous, the 

bracts caducous, the bracteoles deciduous, oblate, 

3-5 mm long, 5-8 mm wide, ciliolate, puberulous 

externally, glabrous within, arising about midway 

on the pedicels; pedicels 7-8 mm long, puberulous, 

the hypanthium inequilateral, 1-2 mm long, ca. 5 

mm wide, the nectaries conspicuous; sepals 18-22 

mm long and 10-12 mm wide, oblong-elliptic or 

oblanceolate-oblong, the margins scarious, puberulous externally, glabrous within; petal red, rosepurple, glabrous, 5-7.5 cm long, 6 cm wide, 

obcuneate-flabelliform, the petalodia 1.5-3 mm 

long, 0.8-1.3 mm wide; stamens all fertile, 22-45 

mm long, 9 of the filaments united basally in an 

inequilateral tube, 12-15 mm long on the ventral 

side, the tube puberulous, the anthers oval, 3.5-5 

mm long and 2-2.5 mm wide; gynoecium glabrous, 

the stigma obtuse, unilateral on the style, the latter 

30-35 mm long, the ovary oblong-obovate, 8-9 mm 

long, 3-4 mm wide, the gynophore 7-9 mm long; 

immature fruits 9-10 cm long, 5-6 cm wide, subquadrate, glabrous, with a bluish bloom when 

immature, the carpophore 14-20 mm long. 

TYPE COLLECTION.-C.F.P. Martius s.n. (hob 

type K, isotypes M), “Para, in sylvis inundatis ad 

insulam Marajo,” Brazil, May 1820. 

DIsTRIBuTIoN.-~ccaSiona~ in the flood forest 

along small rivers or small streams in the area near 

or at the mouth of the Amazon River and in 

French Guiana. Flowering and fruiting April to 

JulyFRENCH GUIANA: No locality or date, Lemee 5.12. (P) 

and Richard s.n. (P). BRAZIL. Terr. Amapa: Rio Apurema, 

22 July 1951, Froes & Black 27607 (IAN, NY, R). Para Mosqueiro, Rio Para, 20 June 1943, Ducke 1260 (NY, R, US); Rio 

Marataua (tributary of Rio Araai), Breves Id., 1 Dec 1922, 

Ducke 16927 (P, RB, U); Mosqueiro, 13 Oct 1923, Ducke 

16928 (RB, U); Belem do Paa, 10 May 1923, Ducke 16929 

(P, RB); Soure (insula Marajo), 13 April 1927, Ducke 20312 

(RB, U, US); Rio Macacoari, Municipio Macapa, 5 July 1951, 

Froes 6 Black 27281 (GH, IAN, NY, US); vicinity of Anapu, 

Rio Cariatuba, Portel, 17 May 1956, Froes 32988 (US); vicinity 

of Anapu, Portel, 10 May 1956, Froes 32776 (IAN), 4 Aug 

1956, Froes 32930 (IAN), 32938 (IAN), 12 Aug 1956, Froes 32962 

(IAN); vicinity of Rio Moju, Rio Cairari, 25 March 1957, 

Froes 33130 (IAN). Gurupa, Martius s.n. (G): Thoe Ass&u

Rio Acara, 27 July 1931, Mexia 5982 (A, BM, F, G, GH, NY, 

U, US); Ilha do Marajo, Rio Paracaruari, 6 Sept 1969, 

Oliveira 4929 (IAN); Belem, IPEAN, Igapo do Catu, 27 May 

1968, Pires C!Y Silva 11758 (IAN) and 3 June 1968, Pires 6 

Silva I1785 (IAN); Colares Island, May 1832, Poeppig 2949 

(NY, W). 

LOCAL NAMES AND UsEs.-Fava do matto (Mexia 

5982); ipe, muirapiranga (Ducke in literature). 

DIscussIoN.-Both E. bijuga and E. glabriflora 

have petals reminiscent of those of E. jenmanii, 

from which both species may have originated, 
NUMBER 28 33 

while retaining the ten fertile stamen condition 

of the “ancestral” E. schomburgkiana. Unlike E. 

glabripora, leaflets of E. bijuga are plane, the inflorescence and sepals are puberulous. The latter 

is typically a tree of the regularly inundated (perhaps even daily?) flood plains along the margins 

of small streams and rivers in the area around the 

mouth of the Amazon; E. glabriflora is a tree of 

the “terra firme,” the land above regular flooding 

in northwestern Brazil, especially near Manaus. 

10. Eperua obtusata Cowan, new species 

FIGURE 11 

Arbor alta, ramulis et foliis glabris; stipulae 

caducae, non visae; petioli 3-4.5 cm longi, valde 

canaliculati, rhachibus 4.5-5.5 cm longis, valde 

canaliculatis; foliola bijugata, petiolulis 4.5-5 mm 

longis, laminis minute pellucido-punctatis, coriaceis, planis, 8.5-11 cm longis, 4-5.5 cm latis, 

elliptico-oblongis, ad basem inaequilateralibus, 

plus minusve rotundato latere inferiore sed acuto 

latere superiore, ad apicem rotundatis, venulis prominulis, nervo marginali distincto; inflorescentiae 

axillares, racemosae, 7.5-9.5 mm longae, axe minute striguloso, bracteis et bracteolis deciduis, ca. 

2 mm longis et 1-2 mm latis, ovatis, acutis, minute 

strigulosis, bracteolis prope basem pedicelli nascentibus, pedicellis 8.5 mm longis, minute strigulosis; hypanthium 2 mm longum, 3.5 mm latum, 

cupulare, minute strigulosum externe; sepala ad 

apicem conspicue glandulosa, angustoelliptica ad 

elliptica, 20 mm longa, 5-8 mm lata, minute strigulosa externe; petalum oblato-obovatum, ad basem attenuatum, ca. 5 cm longum et 6.5 cm latum, 

petalodiis ca. 5 mm longis et 1.5-1.8 mm latis, 

oblongo-lanceolatis; androecium glabrum, 7 staminibus fertilibus, filamentis 11.5-26 mm longis, eis 

fertilium staminum longioribus, 9 in tubo inaequilaterali conjunctis 6.5 mm et 10 mm longo, antheris 3.5 mm longis, 1.5 mm latis, ovalibus, glabris; 

stigma truncatum, stylo 34 mm longo, glabro, 

ovario ca. 5 mm longo et 2 mm lato, obovatooblongo, sericeo, gynophoro 4 mm longo, sericeo; 

fructus non visus. 

TYPE COLLECTION.-LI. Williams 16183 (holotype US sheet 1834953) Capihuara, Rio Casiquiare, Amazonas, Venezuela, 3 Oct 1943. 

DIsTRrBuTIoN.-~nown only from the type COllection. 

DIscussIoN.-h a genus of species characterized 

by acute to acuminate leaflets, the broadly rounded 

ones of E. obtusata are a striking contrast. No 

doubt it is properly referred to this genus but considerable question arises concerning its near relatives. Here I have placed it between E. rubiginosa 

and E. schomburgkiana with which it shares the 

gland-tipped sepals characteristic of the first and the 

inflorescence-type of the second. 

11. Eperua grandiflora (Aublet) Bentham 

FICURE~ 8, 12 

Eperua grandifEora (Aublet) Bentham, F1. Bras. 15(2):226, 

1870. 

Tree 8-30 m tall, the trunk 20-40 cm in diameter, 

the branchlets glabrous, nitid; stipules deciduous 

to persistent, more or less foliaceous, rigid, glabrous, falcate-lanceolate or falcate-elliptic, or inequilaterally oval to elliptic to lanceolate, sometimes lanceolate-triangular, 4-12 mm long, 2-7 mm 

wide; leaves glabrous or sometimes micropuberulous on the upper surface of the base of the costa 

and leaflet blade, the petioles 9-21 mm long, the 

rachis 4.5-12.5 cm long; leaflets 3- or 4-jugate, the 

petiolules 4-8 mm long, the blades epunctate, 

rigid-coriaceous, nitid, plane or marginally revolute, all elliptic to lanceolate or frequently the 

lowermost pair ovate to lanceolate, (4-)7-11.5 

(-15) cm long, 2-5 (-7.3) cm wide, the base of the 

lowest pair or of all but the terminal pair roundedobtuse, the others acute to attenuate-acute, the 

apex acute to longacuminate, mucronulate, the 

venation prominulous to subobscure; inflorescences 

axillary racemes of racemes 3-10 cm long, the axes 

minutely puberulous, the bracts broadly ovate, 2-4 

mm long, 3-4 mm wide, ciliolate, minutely puberulous externally, the bracteoles 3-4 mm long, 

3-5 mm wide, broadly ovate, imbricate, clasping 

the upper part of the pedicel and the hypanthium 

base, pubescent as the bracts, arising at or near 

the pedicel-apex, usually at slightly different 

levels; pedicels 4-5.5 mm long, minutely puberulous, the hypanthium inequilaterally cupular, 1- 

1.5 mm long, 2.5-4 mm in diameter, minutely pu- 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

berulous externally, the nectaries projecting above 

the hypanthial rim; sepals oblong to elliptic, 10- 

12.5 mm long, 2-6 mm wide, minutely puberulous 

externally, the dorsal one concave and larger; petal 

pale pink or lavender-pink, more or less flabelliform, 2.5-3.5 cm long, 2.5-4.5 cm wide, glabrous, 

the petalodia 0.8-2 mm long, 0.4-0.8 mm wide; 4 

or 5 of stamens with larger anthers, the filaments 

of 9 united in an inequilateral tube 3-5 mm long 

on the shorter side, 5-6.5 mm on the longer, the 

filaments 15-30 mm long, puberulous in the united 

region but mostly glabrous above, the anthers oval, 

1.8-2.5 mm long, 1-1.5 mm wide, more or less puberulous or glabrous; stigma obtuse, the style 19- 

28 mm long, the ovary obovate, 3-5 mm long, 

1.5-3 mm wide, densely pilosulose or glabrous, the 

gynophore 3-5 mm long, pilosulose or glabrous; 

fruits not seen. 

11 a. Eperua grandiflora ( Aublet ) Bentham 

guyanensis Cowan, new subspecies 

FIGURE 12 

A ssp. grandiflora foliola marginibus revolutis 

minute venulosis, antheris aliquando leviter minuteque puberulis, ovario gynophoroque dense 

pilosuloso differt. 

TYPE COLLECTION.--T. G. Tutin 245 (holotype 

US sheet 1743474, isotypes BM, K, U), 83 mi 

Bartica-Potaro Road, Guyana, 23 June 1933. 

DIsTRIBuTIoN.-Centra~ to south-central Guyana 

in low forest on white sands. 

GUYANA: Along Berbice-Rupununi Cattle Trail, Berbice 

or Demerara County, 20 May 1919, Abraham 133 (K, NY); 

Essequibo-Mazaruni Divide, 0.5 m S of Bartica, 2 Sept 1932, 

Davis 317 (FD 2310) (K) and 318 (FD 2311) (K); upper 

Rupununi R., near Dadanawa, 24-29 July 1922, de la Cruz 

1704 (GH, NY, US); Moraballi Creek, 11 Nov 1940, Fanshawe 

605 (FD 3341) (K, NY); branch of Eberoabo R., Berbice Co., 

20 May 1919, Hohenkerk 788 (K); Essequibo R., opposite 

Bartica, Nov 1886, Jenman 2482 (K, NY); no precise locality 

or date, Jenman 6269 (K, NY); Great Falls, Demerara R., 

March 1898, Jenman 7304 (K); Essequibo R., Moraballi 

Creek, near Bartica, 20 Sept 1929, Sandwith 314 (K, NY, RB, 

U); no locality, 1844, Rich. Schomburgk 988 (1722) (BM, G, 

GH, K, P, W). 

LOCAL NAMES AND Usm-Ituri wallaba (Sandwith 314); yoboko wallaba (Hokenkerk 788). Used 

for firewood. 

llb. Eperua grandiflora (Aublet) Bentham 

ssp. grandiflora 

FIGURE 12 

Parivoa grandiflora Aublet, P1. Guian. 2:756, pl. 303, 1775. 

Dimorpha grandiflora (Aublet) Willdenow, Sp. PI. 3(2):971, 

Eperua kourouensis Benoist in Lecomte, Not. Syst. 3:273, 7 

1-10 NOV 1802. 

May 1917. 

Leaflets plane; anthers and gynoecium glabrous. 

TYPE COLLECTION.--F. Aublet s.n. (holotype 

BM) “ad ripas fluviorum Guianae,” French 

Guiana. 

DIsTRIBuT1ox.-Frequent in forest along Rio 

Araguari on Brazil-French Guiana Border, coastal 

French Guiana and south-central Suriname. 

SURINAME: Upper Coppename R., Oct 1901, Boon 1209 

(U); Fallawatra, 18 Jan 1972, Iimenez-Saa 1698 (K). FRENCH 

GUIANA: Gourdonville, Kourou R., 8 Sept 1914, Benokt 

1595 (P) (type of E. kourouensis Benoist). BRAZIL. Terr. 

Amapa, Rio Araguari: 2 Sept 1961, Pires et a1 50673 (F, NY, 

R, U, US), 12 Sept 1961, Pires et a1 50878 (GH, NY, RB, U, 

US), 29 Sept 1961, Pires et a1 51307 (G, NY, U, US); vicinity 

of Piedra Fina, 10 Oct 1961, Pires 51659 (F, GH, K, NY, U, 

US). 

DIscussIoN.-This is one of the frequently misinterpreted species in the genus, largely because of 

the specific epithet. For a species characterized by 

a petal among the smallest in the genus, “grandiflora” seems entirely inappropriate. However, compared to the second species of Aublet’s genus Parivoa (a synonym of Eperua), which belongs to 

another genus altogether, the flower is enormous 

so the name must have seemed distinctive. The 

epithet has often been attached to collections of 

E. jenmanii just because of the large flowers of 

the latter. 

It is still a rather poorly known species throughout its range, especially surprising in view of its 

frequency in the boundary area between Brazil 

and French Guiana, as well as in the coastal plain 

of the latter country. It apparently is derived from 

E. jenmanii from which it differs most obviously 

in flower size and petal shape. The two constituent 

subspecies differ in leaflet details, pubescence of 

the ovary, and geographic range. In addition, some 

collections of ssp. guyanensis have flowers whose 

anthers are sparsely puberulous, but this is neither 

very easily observed nor is it always true. 
NUMBER 28 35 

FIGURE lZ.-Geographic distribution of several Eperua species. 

12. Eperua jenmanii Oliver 

FIGURES 8, 12 

Eperua jenrnanii Oliver, Hooker’s Icones 20: 1955, 1891. 

E. stipulata Kleinhoonte, Rec. Trav. Bot. Neerl. 30:171, 1933 

[identity questionable since type is sterile, probably juvenile 

foliage]. 

Medium to large tree 10-35 m tall, the trunk 

15-70 cm in diameter, the branchlets glabrous, 

nitid; stipules glabrous, rigid-foliaceous, free from 

each other or united laterally near their base, 3- 

25 mm long, 1.5-14 mm wide, lanceolate to ovate, 

falcate, acute to acuminate or rounded; petioles 

(8-)14-23 (-45) mm long, glabrous, the rachis 

(4.5-)%I5 (-21) cm long, glabrous; leaflets 3- to 

5-jugate, sometimes the upper ones bijugate, the 

petiolules 4-7 (-12) mm long, glabrous or sparsely 

micropuberulous, the blades pellucid-punctate or 

epunctate, plane to narrowly revolute, coriaceous, 

sometimes thinly so, 6-13 (-18) cm long, 3-8 cm 

wide, the basal pair ovate to lanceolate, the other 

pairs elliptic, the base rounded-obtuse, frequently 

cordulate, the apex acuminate to long-acuminate, 

mucronate, the blades glabrous or micropuberulous 

on the upper surface at the base, the venation 

prominulous; inflorescence a simple axillary or 

terminal raceme, rarely with a few racemose 

branches, the axes 2-6 cm long, glabrous or micropuberulous, the bracts persistent, 2-5 mm long, 

3-6 mm wide, broadly ovate or oval, glabrous or 

minutely puberulous externally, glabrous within, 

often ciliolate, the bracteoles persistent, 3-7 mm 

long, 3-7 mm wide, broadly ovate or oval, clasping 

the upper part of the pedicel and base of the hy- 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

panthium, glabrous or minutely puberulous externally, often ciliolate, arising on the pedicels near 

their base or up to their middle; pedicels 4.5-10 

mm long, sparingly micropuberulous, the hypanthium inequilaterally cupular, 2-3.5 mm long, 5-7 

mm in diameter, glabrous or micropuberulous 

sparingly on the outer surfaces; sepals 13-28 mm 

long, 5-12 mm wide, narrowly oblongelliptic to 

elliptic, elliptic-oblong or oblong, glabrous or micropuberulous externally, the dorsal one broader, 

concave; petal dark lavender to orchid-purple 

with darker purple and white markings, basally 

white, 5-10 cm long, 4-8.5 cm wide, broadly obovate, glabrous, the petalodia 2-10 mm long, linear, 

oblong or triangular-lanceolate; stamens white, 

4-8 apparently fertile (the anthers oblong-oval, 

larger), the filaments of all ten stamens joined 

basally in an inequilateral pilosulose tube 5-1 3.5 

mm long on the shorter side, 8-18 mm long on the 

longer, the dorsal free stamen 32-35 mm long, the 

others 18-65 mm long, most of filaments pilosulose 

basally, glabrous above, the anthers oval, apiculate, 2-4.5 mm long, 1-2.5 mm wide, glabrous; 

gynoecium glabrous, the stigma obtuse to capitellate, the style white, 39-54 mm long, the ovary 5- 

6.5 (-10) mm long, 3-3.5 (-5) mm wide, obovate, 

the gynophore 6-11 mm long; fruits flat, woody, 

oblong to broadly oval, 13-22 cm long, 7-13 cm 

wide, the dorsal margin incrassate, the seeds oval, 

3.5-5.5 cm long, 2-5 cm wide, pale brown or nitidred-brown. 

12a. Eperua jenmanii Oliver ssp. jenmanii 

FIGURES 2c-e and 12 

Stipules 3-9.5 (-20) mm long, 1.5-5 (-10) mm 

wide; leaflets pellucid-punctate; bracts, bracteoles, 

inflorescence axes, and hypanthium minutely or 

microscopically puberulous, the bracts 4-5 mm 

long, 4.5-6 mm wide, the bracteoles 5-7 mm long 

and wide; stamina1 tube 9.5-13.5 mm long on 

shorter side, 12-18 mm on the longer side. 

TYPE COLLECTION.-G. S. Jenman 2154 (lectotype K), Essequibo River, Aug 1884, Guyana, Dec 

1899. In the protologue, Oliver listed five Jenman 

collections of which the one named above is an 

adequate representative. 

DIsTRIBuTIoN.-Frequent to common in lowland 

rain forest, often along rivers, in French Guiana, 

Guyana (on coarse white sands in the Pakaraimas) 

and in southeastern Venezuela at 250-550 m elevation; probably also in Suriname but fertile material from there has not been seen that can be assigned for certain to this species; cultivated in 

Mexico, Singapore, Cameroon (Africa), Jamaica, 

and Trinidad. Flowering August to November. 

VENEZUELA. Bolivar: road south of El Dorado, 42-65 km, 

26 July 1960, Steyermark 86665 (NY, US). GUYANA: Kumaka, 

Kurupung R., 25 Aug 1925, AZtson 313 (K); no locality or 

date, A@un 756 (K): Kartabo Point, Mazaruni R., no date 

or collector, FD 958 (K); Rockstone, 15 July to 1 Aug 1921, 

Gleason 837 (BM, GH, NY, US); Demerara R., Dec 1899, 

Jenman 573 (K, NY), Jan 1882, Jenman 975 (K), Apr 1887, 

Jenman 3830 (K), Sept 1887, 4258 (K); Bartica, Nov 1888, 

Jenman 4770 (K); Kamakusa, 9 Nov 1922, Lang 70 (K, NY, 

US), 9 Nov 1922, Lang G Persaud s.n. (F); Moraballi Creek 

near Bartica, Oct 1929, Sandwith 710 (K, NY); Kukui R., 

Trail from Casobrikeva V, near Adaro R. mouth to Pipilipai 

V, 6 Sept 1960, Tillett G Tilktt 45305 (NY, US); Kako R., 21 

& 25 Sept 1960, TiZZett G Tillett 45474 (GH, NY, US); 

Kamarang R., at mouth of Utschi R., 20 Oct 1960, Tillett Q 

TiZlett 45699 (NY, RB); Cuyuni R., Akarabice Creek, 31 

July 1933, Tutin 439 (BM, K, US); 8.5 miles east of Kaburi 

R., Mazaruni R., 20 April 1926, Wood 876 (K). FRENCH 

GUI.4NA: no locality or date, Martin s.n. [type of Dimor@ 

grandiflora (Aublet) Rudge] (BM). 

LOCAL NAMES AND USES.-Ituri wallaba (Tutin 

439); mahomillo (Steyermark 86665), wobai-yek 

(Altson 313). The wood is used for firewood and 

for shingles. Oliver records the following in the 

protologue: “The scraped root is used by the Indians for the cure of toothache. The timber is used 

for the frames of houses, vat staves, paling staves, 

and shingles for colonial use and exportation.” 

DIscussIoN.-Rudge (Trans. Linn. SOC. 9: 180, 

1808) took up Aublet’s Parivoa grandiflora for 

this species; it is clear from the specimens on 

which Rudge reported his “new species” to the 

Linnean Society, as well as from the description 

and plate, that he grossly misinterpreted Aublet’s 

concept, which he explicitly states he is clarifying. 

The Martin collection is the only record of this 

species in French Guiana and probably came from 

some locality in the interior not collected subsequently. The specimens all show bijugate leaves, 

rather than 3- to 5-jugate ones, but Rudge says that 

the lower leaves are 3- or 4-jugate. There is somewhat more puberulence on the hypanthium/sepals 

but within the range of variation in the species. 
NUMBER 28 37 

12b. Eperua jenmanii Oliver sandwithii Cowan, 

new subspecies 

FIGURE 12 

A var. jenmanii stipulis plerumque 7-25 mm 

longis, 4-14 mm latis, foliolis epunctatis, bracteis 

et bracteoleis inflorescentiarum axibus pediceliis 

hypanthio glabris, bracteis 2-3 mm longis et 3-4 

mm latis, bracteolis 3.5-5.5 mm longis et 3-5 mm 

latis, tubo filamentorum 5-1 1 mm longo dorsualiter sed 8-15 mm longo ventraliter differt. 

otype NY, isotypes F, GH, RB, US), Raudal Guaiquinima, 200 m elev., 16-19 January 1952, Bolivar, 

Venezuela. 

DIsTRIBuTI0N.-Frequent in riverine forest from 

south-central to southeastern Venezuela to Guyana 

and adjacent northern Brazil, at 110 to 1000 m 

elevation. Flowering and fruiting more or less 

simultaneously throughout the year. 

TYPE COLLECTION.-BUSSett Maguire 33136 (holVENEZUELA. Bolivar: Rio Auacapa, 22 Aug 1954, Bernardi 

1536 (NY); between El Dorado and km 88, 9 Sept 1966, 

Blanco 598 (NY, US); Rio Tonoro, Alto Rio Paragua, 2-15 

Aug 1943, Cardona 818 (F, NY, US); mouth of Rio Carapo, 

alto Rio Paragua, 5 Sept 1943, Cardona 985 (US); Rio 

Ikabaru, Oct 1947, Cardona 2158 (US); Rio Paragua, Guaiquinima, 14-15 Apr 1943, Killip 37477 (US); Rio Cuyuni, 

10-15 km below Cerro Escalera, 17 Aug 1962, Maguire et al 

46734 (NY, US); Rio Tirica, lower slopes of Chimana-tepui 

(Torono-tepui) 1000 m elev., 24 May 1953, Steyermark 75537 

(NY); Rio Caroni between mouth of Rio Aparuren & Uriman, 

alt. 400 m, Steyermark 7607# (NY); 3-4 km SE of “Los Patos,” 

30 km S of El Manteco, 9 Aug 1960, Steyermark 87075 (NY); 

Sierra Ichun, along Rio Ichun, 29 Dec 1961, Steyermark 90416 

(US) and 30 Dec 1961, 90458 (NY, US); Rio Paramichi, between mouth of Rio Paramichi & Salto de Halimano, 8-9 

Jan 1962, Steyermark 90724 (NY, US). GUYANA: Demerara 

R., March 1898, Jenman 7305 (K); Membaru-Kurupung Trail, 

2 A’ov 1951, Maguire LT Fanshawe 32435 (NY). BRAZIL. Terr. 

do Roraima: vicinity of Uaica airstrip, Rio Uraricoeira, 7 

March 1971, Prance et a2 10871 (K, NY, U, US). 

LOCAL NAMES AND UsEs.-Concha de suela 

(Blanco 598); opai-yek (Maguire 46734). 

DIscussIoN.-The most conspicuous characteristic of E. jenmanii is its large petal, the largest in 

the genus; only E. bijuga and E. glabriflora compete for this distinction, but both of these have 

only bijugate leaflets and all ten stamens fertile. 

Petal size also separates E. jenmanii from E. grandflora, which are even more closely related. The 

four species form a natural assemblage palynologically, all having unspecialized punctate pollen 

grains. 

The two subspecies are separable, in addition 

to geography, by presence or absence of puberulence on the inflorescence and the size of bracts; 

the stipules tend to be larger and the stamina1 tube 

shorter in subspecies sandwithii. The name of the 

latter commemorates the name of one of the most 

astute legume taxonomists in botanical history. 

His review of the Eperua species that occur in 

Guyana is basic to the present study. 

13. Eperua purpurea Bentham 

FIGURES 2a-b, f-h, 9, and 13 

Eperua purpurea Bentham, F1. Bras. 15(2):226, 1870. 

Large tree (15-)20-50 (-70) m tall, the trunk 

columnar, 30-100 cm in diameter, the bark smooth 

gray, gray-black, or light sand-colored, the branchlets usually glabrous, rarely micropuberulous; stipule more or less fused in one intrapetiolar body, 

persistent to deciduous, broadly semicircular-ovate, 

2-4 mm long and wide, ciliolate, usually glabrous 

externally, more or less sericeous on the inner surface: petioles 10-35 mm long, glabrous, the rachis 

glabrous, 13-80 mm long; leaflets 2- or 3-jugate, 

the petiolules (3.5-)5-7 (-9) mm long, glabrous, 

the blades pellucid-punctate, thinly to rigidly coriaceous, the margins plane or narrowly revolute, 

thicker than the remainder of the blade, glabrous, 

(4.5-)6-10 (-11.5) cm long, 3-6 cm wide, ovate to 

oval or elliptic, sometimes only the basal pair ovate 

and the others elliptic, the base rounded-obtuse, 

sometimes slightly cordate, the apex bluntly acute 

to acuminate, often slightly retuse at the tip, the 

upper surface glabrous, lucid, the lower rarely 

microstrigulose, densely ceriferous except on the 

thickened marginal vein, the wax bodies contiguous, minute, the venation obscure; inflorescence 

a mostly terminal, infrequently axillary, raceme of 

racemes (3-)5-7 (-10) cm long, the axes glabrous 

or rarely micropuberulous, the bracts thin, pellucidpunctate, persistent or deciduous, 1.5-2.5 mm long, 

1-2 mm wide, triangular-ovate, acute, glabrous 

except ciliolate, the bracteoles persistent, similar 

to bracts in texture, pubescence, size, and shape, 

pellucid-punctate, arising at slightly different levels 

on the pedisels about one-third to one-half the 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 13.--Geographic distribution of several Efierua species. 

length from its base; pedicels (9-)lo-I5 (-24) 

mm long, slender, glabrous, somewhat pellucidpunctate, the hypanthium cupular, glabrous, 

pellucid-punctate 2-3 mm long, 2.5-5 mm in diameter, the nectaries usually conspicuous; sepals 

glabrous except more or less ciliolate, pellucidpunctate, elliptic to oval, 9-1 1.5 mm long, 4-9 mm 

wide, the dorsal one broader, concave and cucullate; petal deep rose, pink-purple, purple, rosepurple, 2.5-4.5 cm long, (3.5-)4.5-6.5 cm wide, 

flabelliform, shortly attenuate basally, glabrous, 

the petalodia 0.5-0.7 mm in diameter, semicircular;’ 

stamens 10, 3 to 5 fertile, the filaments of 9 joined 

in an inequilateral villose tube 4-5.5 mm long on 

the shorter side, 5-8 mm on the longer side, the 

filaments of the fertile stamens 18-35 mm long, 

glabrous in the upper half, the sterile ones 12-27 

mm long, villose almost to apex, the one free stamen with the filament 14-15 mm long, villose, the 

fertile anthers oval, 2 mm long, 1.5 mm wide, glabrous; gynoecium glabrous, the stigma capitate or 

capitellate, the style 25-35 mm long, the ovary 3-4 

mm long, 1.5-2 mm wide, obovate, the apex truncate, the gynophore 2-3.5 mm long; fruits 15-17 

cm long, 6-8 cm wide, oblong, flat, with a bluish 

bloom when young, the dorsal margins alate, the 

seeds flat, oval, lucid, red-brown. 

TYPE COLLECTION.--R. Spruce 2577 (holotype 

K, isotypes BR4, BR, F, G, GH, NY, P, RB, W) 

Panure, Rio Negro-Rio Uaupes, Amazonas, Brazil, Sept 1852. 

DISTRIBUTION.-FreqUent to dominant in 

Savanna-scrub, “moist places of the upland rain 

forest” (Ducke, 1940) in the Rio Negro Basin of 
NUMBER 28 

northwestern Brazil, and in similar areas in adjacent southeastern Colombia and southwestern 

Venezuela at 120-750 m elevation. Flowering most 

heavily in October and November; fruiting November to June. 

COLOMBIA. Vaupes: Rio Negro, San Felipe 13-25 Xov 

1952, Hunibert 27386 (US), 24 Oct 1952, Schultes et a1 17955 

(BM, BR, GH, NY, U, US); Rio Guainia, Cano del Caribe 

between Isla del Venado and San Jose, 2 Nov 1952, Schultes 

et al 18268 (GH, U, US); Cerro de Tipiaca, between Mita and 

Javarete on Rio Vaupes, 14-24 May 1953, Schultes &+ Cabrera 

19324 (GH, NY, US). VENEZUELA. Amazonas: Yavita, Rio 

Atabapo, 19 Oct 1950, Maguire 29311 (NY, RB); Playa Alta, 

Rio Cunucunuma, 9 Nov 1950, Maguire et al 29486 (F, NY, 

US); 1 km east of Maroa, 120-140 m elev., 25 Nov 1953, 

Maguire et a1 36398 (NY); just south of Maroa, Rio Guainia, 

28 Nov 1953, Maguire et a1 36452 (GH, NY, U, W); trail to 

Guzman Blanco 1-5 km SE of Maroa, Rio Guainia, 9 Oct 

1957, Magtiire et al 41776 (GH, NY, RB, U, US); Rio 

Guainia, island at Las Isletas, 16 Oct 1957, Maguire et a2 

41922 (F, NY, US); Yavita, 26 Jan 1942, L1. Williams 13990 

(A, F, G, RB, US). BRAZIL. Amazonas: upper Rio Negro, 

June 1932, Ducke 52 (F, NY); Rio Negro above Rio Curicuriary, 19 Nov 1929, Ducke 23288 (G, P, RB, U, US); Rio Negro, 

Ferreira s.n. (P); Jauarete, Vaupes Rio Negro, 19 Oct 1945, 

Froes 21206 (F, IAN, NY, US); Rio Negro, Icana, Santana, 5 

May 1947, Froes 22317 (GH, IAN, U); Rio Cauaburi between 

Cachoeira Jacamin and Manajos, 6 Nov 1965, Maguire et a1 

60112 (F, NY, US); Rio Negro-rio Cauaburi, 27 Nov 1965, 

Magzcire et a1 60421 (NY, US); Rio Negro, Marabitanas, 20 

Apr 1947, Pires 468 (IAN, NY, INPA, US); Rio Uneiuxi, 200- 

300 km above mouth, 22 Oct 1971, Prance et a1 15513 (GH, 

M, NY, U, US); Tapuruquara, 27 Oct 1971, Prance et al 

15609 (GH, M, NY, U, US); Rio Curicuriari, 22-24 Jan 1948, 

Schultes G Lopez 9657-B (GH, IAN, US); Rio Negro, Sao 

Felipe, Serrinha opposite mouth of Rio Issana, 4-7 Apr 1948, 

Schultes LY Lopez 9779 (IAN, R, US); Igarape do Chuva, 

Taracua, Rio Vaupes between Ipanore and confluence with 

Rio Negro, 12 Nov 1947, Schultes c!v Pires 9066 (IAN, US); 

Panure, Rio Vaupes, Feb 1853, Spruce 2577-A (K). 

LOCAL NAMES AND UsEs.-cupaubu-rana (Spruce 

2577), iebaru (Maguire et a1 60112), iobaro (Pires 

468), jebar6 (Spruce 2577), pega-pega (L1. Williams 13990), poo-koo (Schultes et a1 17955, wapa (Schultes et a1 17955), yeba (Schultes et a1 

17955), yebaro (Ducke 23288), yebero (Schultes et a1 

18268), yevaro (Ll. Williams 13990). Wood heavy, 

used for bridges and general construction. Spruce 

records the following information on the holotypelabel: 

This tree, called Jbbaru by the Indians, is the loftiest tree 

in the Caa-tingas and their greatest ornament throughout the 

summer months. It is very frequent in the Rio Negro and 

Uaupes, and when sailing on these rivers its red crown is 

conspicuous from afar, wherever the ground is rising. The 

juice is very viscid and the Indians use it as birdlime for 

taking toucans and parrots. The bark is tough and does not 

decay under water: it is of a red colour. A strip of this bark, 

30 ft. or more long, 11 or 12 inches broad & ’/B inch thick 

is wrapped round a tube, 1 to 3 ft. long, formed of a portion 

of a young trunk of the Paxuiba palm (Iriartea enorrhiza 

Mart.) in a slightly oblique direction, so as to form a widening spiral which extends considerably below the tube, & 

forms a musical instrument which is blown into like a 

trumpet and gives a deep sound that is heard from afar. 

These instruments, called Juruparis, or devils, are in use 

throughout the Uaupes (and anciently it would seem 

throughout the Rio Negro) in the Doboxuris, or feasts, of 

the Indians. I cannot find that they are objects of actual 

adoration, but they certainly are of fear & respect, and several curious observances are connected with them. 

DrscussroN.-Everyone who has seen this species 

in the field, and reported on the experience, has 

exclaimed about the beauty of the tree in flower 

(see Spruce’s description). The inflorescence is 

quite distinctive, partly because of the persistent 

bracts and bracteoles, but the best characteristic to 

separate the species from all other cogeners is the 

ceriferous undersurface of the leaflets. Microscopic, 

wax-secreting glands produce filaments of wax 

which cover the entire lower surfaces except for a 

thickened marginal vein (cf. Figures 1, 3). 

Although the shape and dimensions of the leaflets 

vary considerably, the waxy undersurfaces are always striking, and characters of the inflorescence 

are constant, especially the scarious-margined 

bracts and bracteoles. It is most closely related to 

E. oleifera, which has shorter pedicels and smaller 

nonalate-margined fruits. The pollen grains of 

the two species are very similar and unique in the 

genus by virtue of their reticulate (homobrochate) 

ektexine and broad colpi. 

14. Eperua oleifera Ducke 

FIGURES 9, 13 

Eperua oleifera Ducke, Rev. Bot. Appl. k d’Agric. Trop. 12: 

437, June 1932; Bull. Mus. Nat. Hist. Paris, ser. 2, 4:728, 

OCL 1932. 

Small to large tree, the branchlets glabrous, minutely strigulose or micropuberulous; stipules persistent, joined laterally in a compound, intrapetiolar body 2-4 mm long and 3-4 mm wide, broadly 

ovate, minutely strigulose externally, glabrous 011 

the inner surfaces, ciliolate; petioles 13-23 mni 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

long, usually microscopically puberulous or strigulose, rarely glabrous, the rachis 38-60 mm long, 

pubescent as petioles; leaflets 3-jugate, the petiolules 4-5 mm long, sparingly microstrigulose or 

micropuberulous, the blades pellucid-punctate 

plane, thinly coriaceous to stiffly so, the margins 

usually with a thickened vein, glabrous or rarely 

microstrigulose sparsely below, 5.5-9 cm long, 3.5- 

6.5 cm wide, broadly ovate or oval to elliptic, the 

base rounded-obtuse or cordate, the apex bluntly 

acute to acuminate, the venation prominulous; 

inflorescence a terminal raceme of racemes 3-7.5 

cm long, the axes minutely strigulose to tomentulose, the bracts caducous to deciduous, more or less 

broadly ovate, 1.5-2 mm long and wide, minutely 

strigulose to tomentulose externally, the bracteoles 

persistent at least to anthesis, 1.5-2 inm long and 

wide, broadly ovate to triangular-ovate, pubescent 

as bracts, arising near pedicel-apex; pedicels 4.5-8 

mm long, minutely strigulose to tomentulose, the 

hypanthium 1-2 mm long, 2.5-3.5 mm in diameter, 

equilateral; dorsal sepal larger, concave, oval or 

ovate, the others narrow elliptic and the lateral 

two somewhat arcuate, 7-10 mm long, 2.5-5 mm 

wide, microstrigulose to tomentulose externally, 

glabrous within; petal pale lilac to rose-purple or 

purple, 25-40 mm long, 35-50 mm wide, flabelliform, somewhat attenuate and gibbous basally; 4 

or 5 stamens apparently fertile (the anthers larger), 

the filaments of 9 joined in a strongly inequilateral 

tube 4 mm long on the shorter side and 5-6 mm 

long on the other side, the filaments of the fertile 

stamens 20-30 mm long, the sterile ones and the 

free stamen 12-17 mm long, the tube and the sterile one free, the filaments villose, the anthers 1.5-2 

mm long, 0.8-1 mm wide, oval, those of sterile stamens smaller, malformed, glabrous, acute apically; 

gynoecium glabrous, the stigma obtuse to clavate 

or capitellate, the style 20-24 mm long, the ovary 

obovate, 3-4.5 mm long, 1.5-2 mm wide, the gynophore 3-3.5 mm long; fruit flat, glabrous, ca. 6 cm 

long and 3.5 cm wide, the seed 25 mm long, 18 

mm wide, glabrous, brown-black, nitid, pittedreticulate. 

14a. Eperua oleifera var. oleifera 

FIGURE 13 

Large forest tree with the leaflets oval to elliptic, 

thinly coriaceous, the base rounded-obtuse; petal 

pale lilac. 

TYPE COLLECTION.--A, Ducke 23290 (lectotype 

R, isolectotypes F-frag., G, NY, P, RB, u, Us), 

Borba, Rio Madeira, Amazonas, Brazil, 16 January 

1930. A lectotype choice is required because Ducke 

cited two syntypes in the protologue. 

DIsTRIBuTIoN.-Scattered individuals in noninundated forest along the Rio Madeira and smaller streamlets in southern Amazonia, mostly on 

clay soils. 

BRAZIL. Amazonas: Borba, Rio Madeira, 9 Nov 1935, 

Ducke 73 (US), 5 July 1936, Ducke 213 (A, F, NY, R, US), 

7 Nov 1935, Ducke 235 (NY), 5 July 1936, Ducke 235a (NY). 

LOCAL NAMES AND UsEs.-Jacark copahiba 

(Ducke 232903. Resin from stems used in Brazilian 

Rmazonia in manufacture of paints and varnishes. 

14b. Eperua oleifera Ducke 

var. campestris Ducke 

FIGURE 13 

Eperua oleifera Ducke var. campestris Ducke, Bull. Mus. Nat. 

E. campestris (Ducke) Ducke, Trop. \Voods 62:25, 1940. 

Hist. Paris, ser. 2, 4:728, Oct 1932. 

A small tree of the savannas with the leaflets 

broadly ovate, cordate, more rigid-coriaceous than 

the typical variety; petal rose-purple or purple. 

TYPE COLLECTION.-A. Ducke 23291 (holotype 

R, isotypes G, NY, P, RB, U, US), Borba, Rio 

Madeira, Amazonas, Brazil, 17 January 1930. 

DIsTRIBuTI0N.-Frequent in southern Amazonian savannas, apparently restricted to this area. 

BRAZIL. Amazonas: Borba, Campo Grande, 25 Aug 1942, 

Ducke 73-11 (A, I4N, R, US); Manaus, Sept 1948, Schultes 

6 Lofiez 10337 (GH, IAN, NY, US). 

Ducke made two collections in the Borba area 

which he numbered 73 (9 Nov 1935) and 73-11 

(25 Aug 1942). Apparently the data became confused on the labels during distribution of the duplicates, because 73-11 is clearly variety campestris 

and 73 is the typical variety. The Schultes & Lopez 

collection differs primarily in that the various 

flower parts are minutely tomentulose, rather than 

strigulose as is the material from the type locality. 

DIscussIoh’.-The first publication of the species 

is incomplete and in French but, because it antedates the required Latin diagnosis, it is valid; the 
NUhlBER 28 41 

longer, more complete Latin description later in 

the same year should be consulted to obtain a 

clearer idea of the taxon. Between the two publication dates he discovered a new subspecific taxon, 

variety campestris, which he described briefly in the 

“second protologue,” and later raised it to specific 

level. The author was greatly impressed by the 

difference in stature of the two forms but, at least 

in the herbarium, the differences observed are of no 

more than varietal significance. 

Although there are excellent distinguishing characters to separate E. oleifera and E. purpurea, they 

have a number of common features: (1) nonfoliaceous intrapetiolar stipules; (2) regular hypanthium; (3) fertile stamens reduced to three to 

five; and (4) villose stamina1 tube. They differ 

most markedly in the ceriferous undersurfaces of 

the leaflets in E. purpurea and the glabrous ones 

of E. oleifera. They are scarcely separable palynologically (cf. discussion with E. purpurea). 
Appendix 1: Excluded Names 

Parivoa tomentosa Aublet, P1. Guian. 2:759, 1775. [= Crudia.] 

Eperua hohenkerkii Sprague ex Aitken, Trop. Woods 23:1, 1930. [Nomen 

Eperua decandra Blanco, F1. Filip., ed. 1:368, 1837. [= Afzelia bijuga.] 

Eperua falcata Blanco, FI. Filip., ed. 1:369, 1837. [= Afzelia rhomboidea.] 

nudum.] 

Appendix 2: Lists 

NUMERICAL LIST OF TAXA 

1. E. schomburgkiana Bentham 

2. E. glabra Cowan 

3. E. rubiginosa Miquel 

a. Ex. var. rubiginosa 

b. E.r. var. grandipora Pulle 

4. E. falcata Aublet 

5. E. uenosa Cowan 

6. E. leucantha Bentham 

7. E. duckeana Cowan 

8. E. glabrijlora (Ducke) Cowan 

9. E. bijugu Martius ex Bentham 

10. E. obtusata Cowan 

11. E. grandiflora (Aublet) Bentham 

a. E.g. ssp. guyanensis Cowan 

b. E.g. ssp. grandiflora 

a. E.j. ssp. jenmanii 

b. E.j. ssp. sandzuithii Cowan 

12. E. jenmanii Oliver 

13. E. purfiurea Bentham 

14. E. oleifera Ducke 

a. E.o. var. oleifera 

b. E.o. var. campestris Ducke 

LIST OF EXSICCATAE 

Figures in parentheses refer to the numbers assigned to the 

taxa, which are arranged numerically in the preceding list. 

Abraham, A. A. 

133 (lla) 

Altson, A. H. G. 

313 (12a) 

Aluisio, J. 

172 (7) 

Anderson, C. W. 

122 (4); 182 (1); 183 (3) 

Appun, C. F. 

294 (3); 359 (1); 756 (12) 

Aublet, F. 

s.n. (4); s.n (llb) 

Aubreville, A. 

BAFOG 

236 (4); 260 (4); 313 (3a) 

51M (4); 143M (4); 219M (3a); 1274 (4); 7062 (4); 7064 (4); 

7254 (4); 7298 (4); 7831 (3a); 7937 (3a) 

Bartlett, A. W. 

8150 (4) 

Benoist, R. 

193 (4); 1595 (llb) 

Bernardi, L. 

1536 (12b) 

BGFD 

958 (12a) 

Black, G. A. 

49-8301 (3a); 54-17621 (3a) 

Blanco, C. 

598 (12b); 603 (4) 

Boon, E. 

1199 (4); 1209 (llb) 

Broadway, W. E. 

s.n. (4) 

Burger, D. 

Byron & Elias 

Candido, J. 

s.n. (3a) 

Cardona, F. 

818 (12b); 985 (12b); 2158 (12b) 

Cavalcante, P. 

2538 (3a) 

Coelho, D. 

4308 (7); 5212 (1) 

Crevaux, I. 

s.n. (4) 

Davis, T. A. W. 

317 (lla); 318 (lla); 319 (4); 394 (1); 404 (3a) 

de la Cruz, J. S. 

1373 (4); 1704 (lla); 2091 (1); 2172 (1) 

Dionisio 

4308 (7) 

8 (4) 

67 (8) 

42 
NUMBER 28 43 

Ducke, A. 

14a (8); 52 (13); 73 (14a); 73-11 (14b); 158 (8); 164 (6); 213 

(14a); 235 (14a); 239a (14a); 244 (7); 732 (7); 1260 (9); 9022 

(3a); 16927 (9); 16928 (9); 16929 (9); 20312 (9); 20313 (8); 

23288 (13); 23289 (6); 23290 (14a); 23291 (14b); 23732 (6); 

23734 (7); 23289 (6) 

Elburg, J. 

9822 (4) 

Fanshawe, D. B. 

605 (lla); 644 (1); 2104 (3a) 

Ferreira, A. R. 

s.n. (13) 

Florschiitz, J. & P. A. 

217 (4) 

Focke, H. C. 

845 (4) 

Foldats, E. 

2645 (4) 

Froes, R, L. 

481 (6); 21104 (6); 21206 (13); 22317 (13); 24926 (8); 24949 

(7): 25330 (7); 25698 (3a); 26638 (3a); 26639 (3a); 26649 

(3a); 26687 (4); 32776 (9); 32930 (9); 32938 (9); 32962 (9); 

32988 (9): 33130 (9); 27281 (9); 27607 (9) 

Geay, F. 

s.n. (3a) 

Gleason, H. A. 

332 (1); 412 (1); 837 (12a) 

Goeldi, A. 

1158 (3a): 3862 (8) 

Gonggrijp, J. W. 

2577 (3a); 3700 (3b); 5515 (34 

Guppy, N. G. L. 

198 (3a); 430 (2) 

Halle, F. 

579 (4): 732 (3a) 

Harris, S. A. 

1P-404 (4) 

Hitchcock, A. S. 

17089 (1) 

Hohenkerk, L. S. 

122-B (3a); 788 (lla) 

Hostmann, F. W. 

129 (3h); 229 (3a): 229-A (3b); s.n. (4) 

Hulk, J. F. 

227 (4) 

Humbert, H. 

27386 (13); 27386 bis (6) 

Irn Thurn, E. F. 

INPA 

s.n. (4) 

6065 (8); 6280 (8) 

47471 (3a): 48277 (4); 48408 (4); 48613 (4); 55448 (4); 

55467 (4): 55716 (4); 57579 (3a) 

234 (4); 573 (12a); 627 (1); 768 (3a): 786 (3a); 975 (12a); 

1147 (3a); 1336 (1); 2154 (12a); 2482 (lla); 3830 (12a); 4095 

Irwin, H. S. 

Jenman, G. S. 

(4); 4101 (4); 4258 (12a); 4770 (12a); 4941 (4); 5015 (4); 5257 

(1); 6269 (lla); 7304 (lla); 7305 (12b) 

Jimenez-Saa, H. 

1542 (4); 1597 (4); 1622 (4); 1625 (4); 1627 (4); 1698 (Ilb) 

Kappler, A. 

1689 (3a) 

Kegel, H. 

872 (3a) 

Killip, E. P. 

37477 (12b) 

Krukoff, B. A. 

8006 (8) 

Kunhardt, H. R., Jr. 

Lang, H. 

70 (12a); 276 (4) 

Lang, H., & Persaud, A. C. 

272 (4): s.n. (4) 

Lanjouw, J. 

738 (3a); 1209 (3a) 

Lanjouw, J., & Lindeman, J. C. 

324 (3a) 

Leblond, J. B. 

s.n. (3a) 

Lemee 

s.n. (4); s.n. (9) 

Leprieur, F. 

s.n. (3a) 

Lindeman, J. C. 

4649 (4); 6344 (3a) 

Linder, D. H. 

132 (3a) 

Luetzelburg, P. 

20211 (3a); 20399 (3a) 

Maguire, B., & Maguire, B., et al. 

23804 (3b): 24720 (4); 24778 (4); 29311 (13); 29486 (13); 

33136 (12bj; 36307 (6); 36398 (13); 36452 (13); 36508 (6); 

41776 (13); 41922 (13); 42610 (6); 46734 (12b); 53514 (5): 

53597 (5); 54014 (3a); 54401 (4); 60112 (13); 60130 (6); 60421 

20 (5) 

(1 3) 

Maguire, B., & Fanshawe, D. B. 

32435 (12b) 

Martin, P. S. 

s.n. (4); s.n. (12a); s.n. (3a) 

Martius, P. 

s.n. (9) 

Melinon, M. 

s.n. (4) 

Melo, F. 

Mennega, A. M. W. 

102 (3a); 441 (4) 

Mexia, Y. 

5982 (9) 

Miquel, F. A. W. 

s.n. (4) 

Myers, J. G. 

5672 (3a) 

3913 (7) 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Oliveira, E. 

2737 (7); 4929 (9) 

Osmarino 

s.n. (7) 

Perrottet, G. S. 

23 (3a); s.n. (3a) 

Persaud, A. C. 

14 (1); 166 (1); 243 (4) 

Petrov, I. 

239 (4) 

Pires, J. M. 

468 (13); 531 (6); 978 (6); 48849 (3a); 48876 (4); 50673 (llb); 

50878 (llb); 51182 (3a); 51307 (llb); 51501 (3a); 51659 (llb) 

Pires, J. M. & Silva 

11758 (9) 

Poeppig, E. 

2949 (9) 

Poiteau, A. 

s.n. (34 

Prance, G. T., & Prance, G. T., et al. 

1652 (1); 3072 (7); 3709 (8); 3720 (7); 4637 (8); 4977 (7); 

15513 (13); 15609 (13); 15684 (6) 

Rech 

45 (34 

Richard, L. C. M. 

s.n. (9); s.n. (4) 

Rodrigues, W. 

178 (6); 562 (8); 905 (6); 5756 (8); 8686 (7) 

Rodrigues, W. & Chagas 

1725 (7); 3061 (8) 

Rodrigues, W., Pe Lima, J. 

2233 (7) 

Rodrigues, W. & Osmarino 

5723 (8); 5982 (8); 6769 (7); 6932 (8); 7005 (8) 

Rodrigues, W., et al. 

1807 (7) 

Rombouts, H. E. 

170 (1); 177 (3a); 725 (34 

Sabajo, P. H., & Roberts, L. 

11192 (4) 

Sagot, P. A. 

183 (4) 

Sampaio, A. J. 

5760 (3a); 5818 (34 

Sandwith, N. Y. 

137 (4); 142 (1); 314 (lla); 329 (4): 710 (124 

Schomburgk, Richard 

173 (3a); 174 (3a); 181 (3a); 988/1722 (lla) 

Schomburgk, Robert 

515 (3a); 517 (1) 

Schultes, R. E. 

9066 (13); 9657B (13); 9779 (13): 10337 (14b); 17872 (6): 

17955 (13); 18268 (13); 19324 (13); 19338 (13) 

Schultz, J. P. 

Schweinitz, L. D. 

Smith, A. C. 

Spruce, R. 

Steyermark, J. A. 

7708 (1); 10237 (4) 

s.n. (4) 

2741 (34; 2882 (2) 

2021 (6); 2577 (13); 2577-A (13) 

75537 (12b); 76074 (12b); 86665 (1%); 86692 (4); 87075 (12b); 

90416 (12b); 90458 (12b); 90724 (12b) 

102733 (6) 

311 (3a); 603 (3a); 704 (3a); 806 (1); 1171 (4); 1725 (4); 2577 

(3b); 2750 (4); 3100 (4); 3210 (4); 3385 (4); 3401 (4); 3408 

(4); 3434 (4); 3618 (4); 3625 (4); 3970 (4); 3972 (4); 4061 

(4); 4192 (4); 4200 (4); 4324 (4); 4416 (4); 4420 (4); 4528 

(4); 4536 (4); 4791 (4); 4838 (4); 5042 (4); 5060 (4); 5364 (4); 

5398 (4); 5408 (4); 5420 (4); 5462 (34; 5524 (4); 5547 (4); 

5549 (4); 5804 (4); 5999 (4): 6906 (4) 

45304 (4); 45305 (12a); 45413 (4); 45474 (124; 45699 (12a); 

45791 (4); 45794 (4) 

Tresling, J. 

96 (3a); 102 (3a); 334 (3a) 

Trinidad Herbarium 

1350 (12a) 

Tutin, T. G. 

245 (lla); 439 (12a) 

Vaillant 

s.n. (4); s.n. (3a) 

Versteeg, G. M. 

48 (3b); 157 (3a); 393 (4); 855 (3a) 

Voltz 

s.n. (3a) 

Wachenheim 

130 (4); 405 (4) 

Watson 

Went, F. A. F. C. 

William 

Steyermark, J. A., & Bunting, G. S. 

Suriname Forest Bureau 

Tillett, S. S., & Tillett, C. L. 

58 (1) 

116 (3a) 

501 (7) .. 

Williams, L1. 

13939 (6); 13990 (13); 14390 (6); 14940 (6); 14942 (6); 15801 

(6); 15811 (6); 16183 (10) 

Wood, B. R. 

876 (12a) 

Woodherbarium Suriname 

22 (4); 66 (4) 

Wullschlagel, H. R. 

149 (4); 149-A; (3a); 150 (3a); 2069 (3a); s.n. (4) 
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