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smithsonian contributions to botany • number 109

Understanding Paloue

(Leguminosae: Detarioideae)

Revision of a Predominantly Guiana 

Shield Endemic

Karen M. Redden, 

Patrick S. Herendeen, and 

Gwilym P. Lewis
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smithsonian contributions to botany • number 109

Understanding Paloue

(Leguminosae: Detarioideae)

Revision of a Predominantly Guiana 

Shield Endemic

Karen M. Redden, 

Patrick S. Herendeen, and 

Gwilym P. Lewis

WASHINGTON, D.C.

2018
ABSTRACT

Redden, Karen M., Patrick S. Herendeen, and Gwilym P. Lewis. Understanding Paloue (Leguminosae: 

Detarioideae): Revision of a Predominantly Guiana Shield Endemic. Smithsonian Contributions to Botany, 

number 109, viii + 44 pages, frontispiece + 30 figures, 2 appendices, 2018. — On the basis of morphological 

and molecular phylogenetic analyses, the genus Paloue is revised to include the genera Elizabetha and Paloveopsis. As newly circumscribed, Paloue comprises 17 species, 2 subspecies, and 2 varieties. The following 11 

new combinations in Paloue are made: P. bicolor (Ducke) Redden, P. coccinea (Schomb. ex Benth.) Redden, 

P. duckei (Huber) Redden, P. durissima (Ducke) Redden, P. emarginata (R. S. Cowan) Redden, P. fanshawei (R. S. 

Cowan) Redden, P. leiogyne (R. S. Cowan) Redden, P. macrostachya (Benth.) Redden, P. paraensis (Ducke) 

Redden, P. princeps (M. R. Schomb. ex Benth.) Redden, P. speciosa (Ducke) Redden; and one new hybrid: P. × 

grahamiae (R. S. Cowan) Redden. Based on a total evidence phylogeny, the two varieties of Paloue coccinea are 

no longer recognized, and a lectotype has been designated for Paloue guianensis Aubl. The species of Paloue

are small to large woody trees that are distributed in northern South America; most are endemics in the Guiana 

Shield. All species are taxonomically evaluated and revised based on examination of herbarium specimens, 

alcohol-preserved floral material, rehydrated reproductive material, laboratory studies, label information, and 

field observations. All type material was examined either directly or remotely as scanned specimens. All georeferenced material was used to construct the distribution maps; localities without these data were estimated 

by label locality information, and all were found to occur within the range of the georeferenced species distribution. A key to the species is provided as well as detailed descriptions of all species, subspecies, and varieties. 

Illustrations are included. A list of all specimens examined is provided for each species, and a list of numbered 

exsiccatae is also given.

Cover images, from left to right: Paloue sandwithii inflorescence (detail from Figure 29F); field illustration of 

Paloue × grahamiae (detail from Figure 14); and Paloue riparia inflorescences and developing fruit (detail from 

Figure 30B).

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Library of Congress Cataloging-in-Publication Data

Names: Redden, Karen M., author. | Herendeen, Patrick S. (Patrick Stephen), 1959– author. | Lewis, G. P., 

author. | Smithsonian Institution Scholarly Press, publisher.

Title: Understanding Paloue (Leguminosae: Detarioideae) : revision of a predominantly Guiana Shield 

endemic / Karen M. Redden, Patrick S. Herendeen, and Gwilym P. Lewis.

Other titles: Revision of a predominantly Guiana Shield endemic | Smithsonian contributions to botany ; 

no. 109. 0081-024X

Description: Washington, D.C. : Smithsonian Institution Scholarly Press, 2018. | Series: Smithsonian 

contributions to botany, ISSN 0081-024X ; number 109 | Includes bibliographical references. | 

Smithsonian Institution Compilation copyright 2018

Identifiers: LCCN 2018038586

Subjects: LCSH: Paloue—Guiana Highlands | Legumes—Guiana Highlands.

Classification: LCC QK495.L52 R38 2018 | DDC 583.74—dc23 | SUDOC SI 1.29:109

LC record available at https://lccn.loc.gov/2018038586

ISSN: 1938-2812 (online); 0081-024X (print)

Publication date (online): 29 November 2018

Ó The paper used in this publication meets the minimum requirements of the American National Standard for 

Permanence of Paper for Printed Library Materials Z39.48–1992.
Dedication

We dedicate this manuscript to the memory of 

Claudius Paul Perry (17 November 1977–14 June 2011), 

Guyanese guide and best friend of author Karen Redden. 

Claudius worked with numerous scientists from many 

different disciplines and was an extremely talented 

parataxonomist. Claudius and Karen worked together for 

more than a decade, and he spotted and collected her first 

sample of Paloue in 2003. Without his hard work and 

dedication, this manuscript would not have been possible.

Claudius Paul Perry (17 November 1977–14 June 2011).

Photograph by K. Redden.

Contents

LIST OF FIGURES vii

INTRODUCTION 1

TAXONOMIC HISTORY 2

MATERIALS AND METHODS 4

GENERIC DELIMITATIONS AND AFFINITIES 4

DISTRIBUTION AND HABITAT 5

TAXONOMIC TREATMENT 6

ACKNOWLEDGMENTS 33

APPENDIX A: NUMERICAL LIST OF TAXA 37

APPENDIX B: LIST OF EXSICCATAE 39

REFERENCES 43

Figures

1. Flowers of Paloue, Heterostemon, Elizabetha, and Paloveopsis 2

2. Strict consensus tree showing relationships within the Brownea clade 3

3. Map of the Guiana Shield area of South America 5

4. Geographic distribution of Paloue bicolor 8

5. Geographic distribution of Paloue brasiliensis 9

6. Geographic distribution of Paloue coccinea 10

7. Illustration of Paloue duckei 12

8. Geographic distribution of Paloue duckei 13

9. Illustration of Paloue durissima 14

10. Geographic distribution of Paloue durissima 15

11. Geographic distribution of Paloue emarginata 15

12. Illustration of Paloue fanshawei 16

13. Geographic distribution of Paloue fanshawei 17

14. Field drawing of Paloue × grahamiae 18

15. Geographic distribution of Paloue × grahamiae 19

16. Illustration of Paloue guianensis 20

17. Geographic distribution of Paloue guianensis 21

18. Geographic distribution of Paloue induta 22

19. Illustration of Paloue leiogyne 24

20. Geographic distribution of Paloue leiogyne 25

21. Geographic distribution of Paloue macrostachya 25

22. Geographic distribution of Paloue paraensis 26

23. Geographic distribution of Paloue princeps 27

24. Illustration of Paloue riparia 30

25. Geographic distribution of Paloue riparia 31

26. Illustration of Paloue sandwithii 32

27. Geographic distribution of Paloue sandwithii 33

28. Geographic distribution of Paloue speciosa 33

29. Color photos of Paloue durissima, Paloue macrostachya, 

Paloue brasiliensis, Paloue emarginata, Paloue guianensis, and 

Paloue sandwithii 34

30. Color photos of Paloue induta subsp. induta, Paloue riparia, 

Paloue coccinea, and Paloue × grahamiae 35

INTRODUCTION

Paloue Aubl. belongs to the Brownea clade within subfamily Detarioideae Burmeist. 

(LPWG, 2017; tribe Detarieae of Cowan and Polhill, 1981, and Mackinder, 2005). This 

clade also includes Elizabetha Schomb. ex Benth., Paloveopsis R. S. Cowan, and Heterostemon Desf., genera that are morphologically and geographically similar to Paloue

(Figure 1). Each genus shares a number of morphological characters with at least one of 

the other genera, making generic delimitation and identification problematic. Richard 

Cowan (1957:253) noted this problem and went on to say that “it may be necessary to 

combine some or all of these genera into one.” The results from a recent phylogenetic 

analysis of morphological and molecular data show that Heterostemon is monophyletic and sister to a clade in which Paloue and Paloveopsis are nested within Elizabetha

(Figure 2; Redden et al., 2010). Paloue was first described by Jean Baptise Christophe 

Fusee Aublet in 1775, and the name has priority over the two other generic names. It is 

therefore necessary to combine Elizabetha and Paloveopsis into Paloue. The members of 

Paloue sensu lato are found in northern South America, and many are endemic to the 

Guiana Shield. These plants are medium-sized trees that occur along riverbanks and in 

lowland rainforests. Although the habit and habitat of these plants should facilitate their 

collection, many of the species are represented by only one or a few herbarium sheets. 

Other attributes and life history information of these taxa are also insufficiently studied, 

including animal interactions, pollination biology, and economic importance. The purpose of this investigation is to evaluate the morphological variation among the species 

of Paloue and related genera and produce a revised taxonomic treatment for the group 

based on extensive field observations, herbarium study, and phylogenetic analyses that 

utilize data from morphology and DNA sequences.

Understanding Paloue (Leguminosae: 

Detarioideae): Revision of a Predominantly 

Guiana Shield Endemic

Karen M. Redden,1 Patrick S. Herendeen,2* 

and Gwilym P. Lewis3

1 Formerly with Smithsonian Institution, Department of Botany, National Museum of Natural 

History, MRC 166, Post Office Box 37012, 

Washington, D.C. 20013-7012, USA.

2 Chicago Botanic Garden, 1000 Lake Cook 

Road, Glencoe, Illinois 60022, USA.

3 Comparative Plant and Fungal Biology Department, Royal Botanic Gardens, Kew, Richmond, 

Surrey, TW9 3AB, UK.

* Correspondence: pherendeen@chicagobotanic.

org

Manuscript received 17 October 2011; resubmitted 1 May 2018; accepted 11 June 2018.
2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

TAXONOMIC HISTORY

Aublet (1775) described and named the genus Paloue. 

He learned of the name from the indigenous people of French 

Guiana (Aublet, 1775). According to Jussieu (1789), Aublet’s 

spelling of Paloue was not correct and he renamed the genus 

Palovea (orthographical variant, orth. var.). Palovea was widely 

accepted and numerous publications cited Jussieu’s interpretation of the spelling of the genus name. In 1906 Pulle described 

a new species of Palovea and named it P. riparia Pulle based on 

the specimen he collected on the banks of the Saramacca River, 

near Janbasigado in what was then called British Guiana (Pulle 

FIGURE 1. Flowers of Paloue (a), Heterostemon (b), Elizabetha (c), and Paloveopsis (d). Reproduced from Cowan (1957) with permission from New York 

Botanical Garden.

1906), now Guyana. Ducke (1915) described Palovea brasiliensis from the forest near the Cachoeira do Apuhy on the Tapajoz 

River in Brazil. He also used Jussieu’s name, Palovea. Sandwith 

(1937) described the most recently published species, Paloue

induta Sandwith, with the genus name reverting back to Aublet’s 

orginal spelling. Scopoli (1777) used the same specimen as Aublet (1775) to describe his new genus, Ginannia Scop. This name 

was quickly recognized as a synonym of Paloue and was subsequently used in the literature only four times. Necker (1790) cited 

Scopoli but spelled the genus name Ginnania, which resulted in 

further confusion. All recent publications that recognize Paloue

use this spelling and attribute the authorship to Aublet.
Schotia spp.

Barnebydendron riedelii

Saraca declinata

Cynometra megalophylla

Macrolobium bifolium

Macrolobium campestre

Macrolobium gracile

Mlacrolobium longipes

Macrolobium multijugum var. multijugum

Macrolobium sp. 1

Ecuadendron acosta-solisianum

Brachycylix vageleri

Brownea coccinea

Paloue bicolor var. velutina

Paloue coccinea var. coccinea 1

Paloue coccinea var. coccinea 2

Paloue coccinea var. coccinea 4

Paloue coccinea var. coccinea 3

Paloue duckei

Paloue durissima 1

Paloue durissima 2

Paloue durissima 3

Paloue leiogyne

Paloue macrostachya

Paloue paraensis 1

Paloue paraensis 2

Paloue princeps 1

Paloue princeps 2

Paloue speciosa

Heterostemon conjugatus

Heterostemon ellipticus

Heterostemon impar

Heterostemon ingifolius

Heterostemon mazarunensis

Heterostemon mimosoides var. complanatus

Heterostemon mimosoides var. mimosoides

Heterostemon mimosoides var. pacimoniensis

Heterostemon otophorus 2

Heterostemon otophorus 1

Paloue emarginata 2

Paloue emarginata 1

Paloue brasiliensis

Paloue guianensis 2

Paloue guianensis 4

Paloue guianensis 5

Paloue guianensis 3

Paloue guianensis 1

Paloue induta 2

Paloue induta 1

Paloue induta 5

Paloue induta 4

Paloue induta ssp. glabra

Paloue riparia 2

Paloue riparia 1

Paloue sandwithii 2

Paloue sandwithii 1

Paloue induta 3

Paloue riparia 3

100

98

66

63

79

60

100

91

100

66

50

99

74

98

56

87

86

64

96 100

93

100

97

81

78

98

100

55

100

58

61

100

98

68

73

67

61

100

95

100

86

67 50

51

99

74

96

60

90

89

68

87

100

66

82

97

96 100

100

100

96

54

98

57

79

Brownea

 Clade

FIGURE 2. Strict consensus of the 234 shortest 

trees based on parsimony analysis of the Brownea

clade total evidence data set (morphology, ITS, 

and trnL). Bootstrap and jackknife values > 50% 

are above and below the nodes, respectively 

(1,476 characters, L = 986, C.I. = 45, R.I. = 0.74). 

From Redden et al. (2010).
4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

The genera Elizabetha and Paloveopsis have straightforward nomenclatural histories. Elizabetha was described by Bentham (1840) and Paloveopsis by Cowan (1957). Cowan’s (1976) 

revision of Elizabetha included all recognized species, and since 

its publication no new species have been described.

MATERIALS AND METHODS

All species were directly observed from herbarium sheets 

(BM, BRG, F, K, L, MO, NY, P, U, US, WAG; Thiers, continuously updated) from the following institutions:

BM Herbarium, Algae, Fungi and Plants Division, The 

Natural History Museum, London, UK

BRG Guyana National Herbarium, Biology Department, 

University of Guyana, Georgetown, Guyana

F Herbarium, Botany Department, Field Museum of 

Natural History, Chicago, Illinois, USA

K Herbarium, Royal Botanic Gardens, Kew, Richmond, 

Surrey, UK

L National Herbarium Nederland, Botany Section, 

Naturalis Biodiversity Center, Leiden, Netherlands

MO Herbarium, Missouri Botanical Garden, Saint Louis, 

Missouri, USA

NY William and Lynda Steere Herbarium, The New York 

Botanical Garden, Bronx, New York, USA

P Herbier National, Muséum National d’Histoire 

Naturelle, Paris, France

U National Herbarium Nederland, Botany Section, 

Naturalis Biodiversity Center, Leiden, Netherlands

US United States National Herbarium, Department of 

Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA

WAG National Herbarium Nederland, Botany Section, 

Naturalis Biodiversity Center, Leiden, Netherlands

Other abbreviations used in synonymies and specimens 

examined sections or elsewhere in text are defined as follows:

comb. nov. new combination

fl. flower

fr. fruit

imm. fr. immature fruit

s.n. sine numero (unnumbered collection)

In addition, field observations, measurements, and notes 

were integrated into the descriptions. In general, vegetative measurements were made from dried herbarium material, whereas 

reproductive structures were measured directly from field specimens, alcohol-preserved specimens, and rehydrated herbarium 

specimens. Length of leaf and leaflet is given from the apex of 

the leaf or leaflet lamina to the basalmost insertion point of the 

blade on the petiole or petiolule. The petiole length is also measured from this point. Colors of corollas, fruits, and other organs 

are described from herbarium label data, field observations, 

and original descriptions. Conventions used in measuring and 

describing various organs follow Cowan’s revision of Elizabetha

(Cowan, 1976). In his revision, Cowan refers to the supporting 

stalk of the hypanthium as the “stipe,” which is measured from 

the insertion of the bracteoles to the base of the hypanthium. 

Collection locality details in the specimens examined sections are 

taken directly from herbarium labels and given verbatim without 

translation.

Cowan’s (1976) revision of Elizabetha was complete and comprehensive, and this monograph follows his convention for all taxa 

now included in Paloue. Much of the taxonomy for Elizabetha and 

Paloveopsis was taken directly from his revision with minor modifications to measurements, additional information about structures 

not previously seen, new locality information, and addition of 

recent collections.

GENERIC DELIMITATIONS AND AFFINITIES

Paloue historically has been placed in the Brownea group 

of the tribe Detarieae within the subfamily Caesalpinioideae 

(Cowan and Polhill, 1981). Other members of this group have 

included Barnebydendron J. H. Kirkbr. (synonym: Phyllocarpus

Riedel ex Tul.), Eperua Aubl., Ecuadendron D. A. Neill, Paloveopsis, Elizabetha, Brachycylix (Harms) R. S. Cowan, Heterostemon, Brownea Jacq., and Browneopsis Huber. The Brownea 

group is now part of the recently reinstated subfamily Detarioideae (LPWG, 2017), but the genera traditionally placed in the 

Brownea group are considered to belong to three different tribes 

(de la Estrella et al., 2018), with Eperua and Barnebydendron

not grouping with the other genera retained within it. The group 

also now includes the diverse neotropical genus Macrolobium (de 

la Estrella et al., 2018; Murphy et al., 2018; Schley et al., 2018). 

In Cowan’s (1957) description of Paloveopsis, he compared the 

morphological attributes of many of these genera and noted that 

each shares a number of common characters with the other genera, leading to a blurring of generic boundaries (Cowan, 1957; 

Figure 1). A phylogenetic analysis of the tribes Detarieae and 

Amherstieae using chloroplast trnL intron sequences has shown 

that these two tribes together form a monophyletic group but 

that neither tribe individually is monophyletic (Bruneau et al., 

2000). An analysis of the Brownea clade using 125 morphological, anatomical, and reproductive characters grouped Paloue, 

Paloveopsis, Elizabetha, and Heterostemon into a monophyletic 

group (Redden and Herendeen, 2006). However, support for 

nodes along the backbone of the tree was low. In a recent phylogenetic study using combined morphological and molecular data 

[chloroplast trnL spacer and nuclear internal transcribed spacer 

(ITS)], it was shown that Elizabetha, Paloveopsis, and Paloue

together form a monophyletic group that is sister to a monophyletic Heterostemon (Redden et al., 2010; Figure 2). These results 

have been corroborated by Murphy et al. (2018) and Schley et al. 

(2018). Nevertheless, it is apparent that there is a distinct difference between the white-flowered Paloue (formerly Elizabetha, 

in part) and the red-flowered Paloue. The size of the flowers, 

leaf structure, size of the leaflets, and pollen type differ. A few of 
NUMBER 109 • 5

the red-flowered species are known to hybridize, something not 

observed in the white-flowered species. These differences should 

be investigated further in future studies. The monographic revision presented here is based on the total evidence phylogeny 

obtained in the study of Redden and Herendeen (2006).

Within the Brownea clade, Paloue s.l. can be distinguished 

from other members by a unique combination of morphological characters. The leaves range from simple to multijugate. In a 

number of specimens of P. guianensis a pair of lateral, vestigial 

leaflets has been noted at the base of the “leaf,” suggesting that 

the “simple” leaves are actually unifoliolate and a reduction from 

a compound leaf. These vestigial leaflets are caducous and have 

only been seen in specimens that included very young leaves. The 

androecium of Paloue consists of either nine fertile stamens, or 

three fertile stamens and six staminodes with vestigial anthers 

(as in P. grahamiae), or three fertile stamens plus six staminodes 

without vestigial anthers. The fruit is a flattened pod, usually 

with a lip or a flange along the placental margin, a character not 

found in its closest relative, Heterostemon. The pollen exine is 

gemmate to verrucate, unlike the striate pollen in Heterostemon.

DISTRIBUTION AND HABITAT

Species of Paloue are predominately found in riparian and 

lowland rainforests of Brazil, Venezuela, Suriname, French Guiana, and Guyana and are almost exclusively concentrated in the 

Guiana Shield (Figure 3). One species, Paloue duckei (Huber) 

Redden, is found in Colombia. Many of these species are represented by only a handful of specimens, and as collection efforts 

FIGURE 3. Map of the Guiana Shield area of South America. Map by Thomas H. Hollowell, Smithsonian Institution.
6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

increase, knowledge of distributional patterns may change and 

geographic ranges increase. Most species are found at elevations 

below 500 m, but a few occur predominantly in the foothills 

of mountain ranges and in the tepui regions of Venezuela and 

Guyana. Most species are found on poor soils or white sands in 

seasonally flooded forests. There exist few data on nodulation or 

mycorrhizal fungi association for these species. Five species of 

Paloue (as Elizabetha) have been reported as non-nodulating and 

one (E. coccinea) as nodulating (Souza et al., 1994), although the 

positive report needs to be checked (Sprent, 2001). There are no 

reports on nodulation for Paloue sensu stricto and Paloveopsis

(Sprent, 2001).

TAXONOMIC TREATMENT

Paloue Aubl., Historie Des Plantes de La Guiane Francoise. 

Paris, Ches Pierre-Francoise Didot. 1775. Type: P. guianensis Aubl.

Elizabetha Schomb. ex Benth., J. Bot. (Hooker) 2: 92. Mar 1840. Lectotype: 

E. princeps Schomb. ex Benth. (R. S. Cowan, Taxon 8: 59. 1959).

Paloveopsis R. S. Cowan, Brittonia 8: 251. 9 Jan 1957. Type: Paloveopsis 

emarginata R. S. Cowan.

Small to large trees, 7–50 m tall, branchlets brown to gray, 

glabrous to pubescent. Stipules intra-petiolar, persistent to caducous, lanceolate to oblanceolate–obovate, linear to foliaceous, 

thin to membranous, joined on inner margin throughout or 

shortly above the base, sometimes bifid. Leaves simple to multijugate, usually paripinnate but imparipinnate in one species 

(P. grahamiae), alternate; petioles 0.5–5 mm long; when leaves 

simple (unifoliolate), blade margin entire, oblong to lanceolate 

to ovate, coriaceous, pinnately nerved, glabrous, apex obtuse 

to long-acuminate, base rounded to acute to auriculate; when 

multijugate, 2–57 leaflet pairs, the leaflets narrowly oblong to 

elliptic; rachis usually narrowly alate, the wing interrupted at 

insertion of leaflet pairs, often with short distal extension of the 

wing at junction of leaflet pairs. Inflorescences dense or open, 

spike-like, terminal or axillary racemes, bracts orbiculate, caducous or persistent, bracteoles connate into a bilobed tube enclosing the flower bud, or shorter than the calyx, rounded, persistent 

beyond anthesis; pedicels 2–22 mm long, hypanthium 10–29 

mm deep including the stipe, glabrous to pubescent. Sepals 4, 

usually unequal in shape with the dorsal sepal broader, reflexed, 

glabrous to pubescent. Petals 5, subequal or of different sizes. 

Fertile stamens 3 or 9, filaments of fertile stamens 2–6 cm long, 

connate with staminodes basally to form a tube or connate 

except at dorsal petal, glabrous to pilose, anther oblong, uniform 

(except in P. grahamiae), glabrous to densely pilose. Stigma capitate, style elongate, filiform, the ovary narrowly oblong, velutinous to glabrous, the stipe shorter than the ovary, adnate to the 

upper surface of the hypanthium tube, ovules numerous. Fruit 

a legume, straight to curved, oblong to scimitar-shaped, twovalved, compressed, woody, the upper sutures thickened with a 

ridge or lip perpendicular to the valve face or parallel to the pod 

surface (P. coccinea); seeds ovate, compressed.

Key to the Species of Paloue

1. Leaves simple, usually with a pair of crater-like glands at the base of the lamina, one on either side of the main vein . . . . 2

Leaves compound, crater-like glands present but usually scattered on lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

2. Ovary glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Ovary pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

3. Staminodes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. P. emarginata

Staminodes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Stamen filaments less than 2.5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. brasiliensis

Stamen filaments greater than 2.5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Petals approximately equal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. P. sandwithii

Petals not equal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Petals: 3 large (2–3 cm long) plus 2 small (2–3 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. P. guianensis

Petals: 1 large (5–7 mm long) plus 4 small (1–3 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. P. riparia

7. Petals: 3 large (2–3 cm long) plus 2 small (2–3 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. P. guianensis

Petals: 1 large (1.3–1.4 cm long) plus 4 small (1.8–3.5 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Inflorescence covered with golden brown trichomes; densely tomentose . . . . . . . . . . . . . . . . 10a. P. induta subsp. induta

Inflorescence sparsely tomentose to glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10b. P. induta subsp. glabra

9. Leaves 2–12 jugate; pedicels mostly 1 cm long or longer, the hypanthium glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Leaves 18–57 jugate; pedicels always less than 1 cm long, usually much less, the hypanthium glabrous or pubescent . . . . 13

10. Leaves imparipinnate, the terminal leaflet larger than lateral leaflets, with paired glands at base of lamina . . . 8. P. × grahamiae

Leaves paripinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Inflorescences 10–100 cm long; petals red or white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Inflorescence to 8 cm long; petals white, the dorsal one well developed and the other 4 reduced to petaloidia about 2 mm 

long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. P. fanshawei
NUMBER 109 • 7

12. Leaflet base equal, acute; 3 fully developed petals and 2 reduced, usually white; staminodes absent; fruit with woody lip 

perpendicular to face of legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. P. macrostachya

Leaflet base unequal, obtuse; 5 fully developed petals, usually red; staminodes present; fruit with woody lip parallel to face 

of legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. coccinea

13. Leaf rachis wing with apical extensions at each pair of leaflets [see Figure 10C] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Leaf rachis wing without apical extensions at each pair of leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

14. Flowers white; hypanthium pubescent; petals dimorphic, the dorsal usually larger and broader than laterals . . . . . . . . 15

Flowers red; hypanthium glabrous; petals about equal in size and shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

15. Inflorescence dense, spike-like; hypanthium tube shorter than its stipe; ovary velutinous; leaflet apex obtuse or truncate . . . 16

Inflorescence elongate, not dense, nor spike-like; hypanthium tube longer than its stipe; ovary glabrous; leaflet apex 

acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. P. durissima

16. Leaflets about 6 or 7 times as long as wide, tapering to a rounded tip; bracts mostly 28–38 mm long and 24–35 mm wide, 

the bracteoles 20–23 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. P. princeps

Leaflets broader, apex usually truncate or rounded, often retuse, rarely tapering apically; bracts 15–27 mm long and 8–15 

mm wide, the bracteoles 14–19 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. P. paraensis

17. Fertile stamens 37–63 mm long; bracteoles united for a greater length on one side; ovary glabrous; leaflets rounded 

apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. P. duckei

Fertile stamens 19–22 mm long; bracteoles equally united; ovary velutinous; leaflets truncate apically . . . 17. P. speciosa

18. Leaves mostly 37–50 jugate, leaflets 19–28 mm long, the undersides with a tuft of long hairs on one side of the main vein 

at its base; inflorescence 50–70 mm long; hypanthium glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. P. leiogyne

Leaves 23–34 jugate, leaflets 11–16 mm long; hypanthium strigulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Ovary pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. P. bicolor var. velutina

Ovary glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. P. bicolor var. bicolor

1. Paloue bicolor (Ducke) Redden comb. nov.

Elizabetha bicolor Ducke, Trop. Woods 1934. No. 37: 22. 1 Mar 1934. 

Type. Brazil. Pará: Rio Itapacura, affluent Rio Tapajoz, Cachoeira 

do Americano, 25 Jan 1933 (fl., fr.). A. Ducke 23726 (RB # 34-961)

(holotype: RB; isotypes: G, K!, P!, U!, US!).

FIGURE 4

Small tree 6–15 m tall, the trunk 10–20 cm in diameter, 

branchlets brown to gray, pilose or pilosulose, hairs dark and 

shiny. Stipules caducous, membranous, narrowly oblong–oblanceolate, apex slightly bifid, pilosulose externally at the base but 

otherwise glabrous. Leaves compound, alternate, paripinnate; 

petioles 1–2.5 mm long, pilosulose; rachis 6–11 cm long, the 

rachis wing narrow and spreading without apical projections, 

contracted at each leaflet insertion, the axis pilosulose; leaflets

11–16 × 2–3 mm, 23–34(–38)-jugate, sessile, narrowly oblong to 

lanceolate–oblong, the base obtuse, inequilateral, apex rounded 

and minutely apiculate, glabrous, the lower surface microscopically red-dotted. Inflorescences 2.5–7 cm long, dense racemes, 

axillary, or terminal and axillary, at tips of branchlets, usually 

densely strigulose to strigose; bracts 10–17 × 14–21 mm, caducous, oblate to sub-rotund, red, strigulose both inside and out, 

margins ciliolate; pedicels 2–6 mm long, puberulous or strigulose; bracteoles 9–17 × 8–9 mm, caducous, connate, united marginally in a tube enclosing the flower bud, rounded, more or less 

carinate dorsally, red, strigulose inside and out, or glabrous on 

inner surfaces. Flowers: hypanthium red or white, strigulose; 

stipe 2.5–4(–9) mm long; tube 7.5–12 × 5–7.5 mm, cylindrical; 

sepals 11–15 × 5–9 mm, oval–oblong, the dorsal one larger, red 

or white, strigulose externally but glabrous on inner surfaces, the 

margins ciliolate; the dorsal petal 10–18 × 8–10 mm, ovate, cordate–auriculate basally, emarginate apically, the other four petals 

12–14 × 5–7 mm, obovate–oblanceolate, red or white, ciliate; 

androecium monadelphous forming a sheath, 2.5–3 mm deep, 

open at insertion of dorsal petal; fertile stamens 3, white; filaments 18–23 mm long, villose on lower parts; anthers 5–6.5 × 

1.5–2 mm, narrowly oblong, glabrous; staminodia 6, 4.5–6 mm 

long; gynoecium dark red; ovary 7.5–12 × 2–2.5 mm, narrowly 

elliptic to elliptic–oblong, velutinous or glabrous; gynophore 

2–8 mm long, adnate to upper surface of the hypanthium tube 

near its rim, geniculate–arcuate, glabrous or puberulous near the 

apex, style 12–23 mm long, glabrous, stigma capitellate. Immature fruit 12–22 × 3–4.5 cm, the dorsal suture bi-marginate, velvety or glabrous, ca. 6–8-seeded.

1a. P. bicolor (Ducke) Redden var. bicolor

Ovary glabrous.

Distribution. Brazil: Amazonas and Pará.

Phenology. Known to flower and fruit in January.

1b. P. bicolor (Ducke) Redden var. velutina (R. S. Cowan) Redden, comb. nov.

Elizabetha bicolor Ducke var. velutina R. S. Cowan, Proc. Kon. Ned. 

Akad. Wetensch. Ser. C. 79: 334. 1976. Type. Brazil. Amazonas: 

Parintins, above Lake Jose-Assu, 29 Dec 1935 (fl.), A. Ducke 113

(holotype: RB; isotypes; F!, G, K!, NY!, P!, R, RB, U!, US!).

Ovary velutinous.
8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

5 Mar 1970 (fr.), Rodrigues 8730 (US!). Pará: Rio Tapajoz collines des environs des cataractes du Mangabal, 10 Feb 1917 

(fl., fr.), Ducke 16751 (US!); 10 Dec 1919 (fr.), Ducke s.n. (RB 

10984, K, 2 sheets!).

2. Paloue brasiliensis Ducke, Arch. Jard. Bot. Rio de Janeiro 

1: 27. 1915. Type. Brazil. Pará: Habitat ad fluvium Tapajoz in silvis ad Cachoeira do Apuhy 1, 28 Dec 1908 (fl., 

fr.), E. Snethlage s.n., H.A.M.P. 10.117 (holotype MG; 

isotype: F!).

FIGURES 5, 29C

Small to medium-sized tree 12–25 m tall, branchlets gray/

brown, glabrate. Stipules caducous, not seen. Leaves simple, alternate, entire, 8.5–16 × 3–7 cm, the blade oblong–

lanceolate, glabrous, coriaceous; petiole 3.5 mm long, base 

rounded to truncate, usually with paired crater-like glands at 

the base of the lamina, apex long-acuminate. Inflorescences

11–14 cm long, elongated terminal racemes; pedicels 6–10 mm 

long; bracts 10 mm long, caducous, orbiculate; bracteoles persistent beyond anthesis, connate, bilobed, carinate, rounded, 

glabrous. Flowers: hypanthium 29 mm long, glabrous, tubular, 

red; sepals 4, 1.1–1.5 × 0.7 cm, reflexed, rounded at the apex, 

dark red; petals 5, largest petal 20–22 × 6 mm, clawed, second petal 6–18 × 4 mm, third petal 8 × 8 mm, the remaining 

two petals are minute, red, margins of all petals pilose; stamens 9, red; filaments 2–2.5 cm long, connate to form a tube, 

sparsely long-pilose; anthers 5–7 mm long, oblong, sparsely 

pilose; ovary 10 mm long, narrowly elliptic to elliptic–oblong, 

glabrous, stipe adnate to upper surface of the hypanthium tube 

near its rim; style 2 cm long; stigma capitate. Fruits 11 × 3 cm, 

oblong, margin slender, stipitate, transversely veined, acuminate, glabrous, ca. 4–8-seeded.

Distribution. French Guiana; Brazil: Pará.

Phenology. Flowering from August to December; 

fruiting from February to May and in October.

Additional specimens examined. BRAZIL. Pará:

Rio Tapajoz, cachoeiras inferiors, lugar Pimentel, 28 Aug 1916 

(fl., fr.), Ducke s.n. (MG-16416, RB 10882, K!, U!, US!); Parque 

Nacional do Tapajos, km 60 da estrada Itaituba–Jacarecanga; 

ilha em frente a Pimental, 26 Sep 1978 (fr.), Silva & Rosario 

3967 (MG, INPA, K!, US!); Rio Tapajoz (near Pará) loco Bella 

Vista, 6 Dec 1919 (fl.), Ducke s.n. (RB 10883, BM!, INPA, K!, 

U!, US!).

FRENCH GUIANA. Cayenne: Rives de l’Oyapock á la hauteur des Itets Yacarescin, 10 Dec 1965 (fl.), Oldeman 1734 (P!, 

U!, US!). Campoi: Rivière Yaroupi, rive gauche, environ 9 km 

en amont du Saut Mouroucioutou, 18 Apr 1970 (fr.), Oldeman

3030 (CAY!, K!); Rives du bas de la crique Inipi (affluent de 

la rivière Camopi), 3 km de son embouchure, 7 Feb 1968 (fr.), 

Oldeman & Sastre 77 (P!, U!, US!); Rives du Camopi (affluent de l’Oyapock) entre les sauts Petit Impossible et Grand 

Impossible, 10 Feb 1968 (fl.), Oldeman & Sastre 163 (U!, US!). 

FIGURE 4. Geographic distribution of Paloue bicolor (Ducke) 

Redden subsp. bicolor (dots) and P. bicolor subsp. velutina (R. S. 

Cowan) Redden (triangles) in Suriname and Brazil.

Distribution. Brazil: Amazonas and Pará.

Phenology. Flowering from November to February 

and fruiting from December to March.

Additional specimens examined. BRAZIL. Amazonas: Borba, Rio Madeira inferior near Amazonas, 16 Jan 

1930 (imm. fr.), Ducke s.n. (RB 23287, K!, US!); Rio Itapecerica 

affluent Tapajoz near Pará, 25 Jan 1953 (fl.), Ducke s.n. (RB 

23727, K!, US!); Borba, Rio Madeira (near Amazonas), 8 Nov 

1935 (fl.), Ducke s.n. (RB 34-959, K!, US!); Parintins (Amazonas), 29 Dec 1935 (fl.), Ducke s.n. (RB 34-960, K!, U!, US!); 29 

Dec 1935 (fl.), Ducke s.n. (RB 34-961, K!); Mt. Dourado, Gleba 

Pacanari, 0°40′S, 52°35′W, 20 Jan 1988 (fl.), Pires & Silva 1954

(K!, NY!); Monte Dourado, Estação Ecol. do Jari (SEMA), 

0°27′S, 52°51′W, 4 Feb 1988 (fl.), Pires & Silva 1984 (K!, 

NY!). Manaus–Caracarai km 60, Reserva INPA-SUFRAMA, 

9 Dec 1976 (fl.), Prance & Ramos 24350 (A, C, COL!, F!, 

INPA, K!, M, MG, MICH, MO!, NY!, P, R, S, U!, UFMG, US!, 

VEN!); Estrada Manaus–Itacoatiara, km 68, 29 Feb 1968 (fr.), 

Rodrigues et al. 8461 (US!); Estrada Manaus–Itacoatiara, km 68, 
NUMBER 109 • 9

FIGURE 5. Geographic distribution of Paloue brasiliensis Ducke 

(dots) in French Guiana and Brazil.

Roura: Bas-Camopi, 21 May 1965 (fr.), Cremers 1328 (CAY!); 

Rivière Counana, 5 km en aval de Dégrad Lalanne, 4°32′N, 

52°17′W, 6 Oct 2000 (fr.), Sabatier & Prévost 4674 (CAY!, 

US!). St. Georges: Fleuve Oyapock, entre St. Georges et 

Camopi, 3°38′25′′N, 51°59′42′′W, 25 Feb 2000 (fr.), Fleury & 

Pignal 1691 (CAY!, US!). Saint-Laurent du Maroni: Marouini 

River, Ile d’Antecume Pata, confluent de l’Itany (Haut Maroni) 

et du Marouini, 17 Nov 1977 (fl.), Cremers 5063 (CAY!, P!, 

U!, US!).

3. Paloue coccinea (M. R. Schomb. ex Benth.) Redden comb. 

nov.

Elizabetha coccinea M. R. Schomb. ex Benth., Hook. Jour. Bot. 292. 1840. 

Type. Guyana: without locality but probably Upper Essequibo River 

(Cowan, 1976), R. Schomburgk s.n. (holotype: K!).

Elizabetha coccinea var. oxyphylla (Harms) R. S. Cowan, Proc. Kon. Ned. 

Akad. Wetensch. 79: 340. 1976. Type. Brazil. Rio Branco, Gebiet des 

Surumu, Ufer des Yalburg, Nov 1909 (fl., fr.), E. Ule 8146 (holotype: B, 

destroyed; isotypes: F!, G, K!, L,! RB).

FIGURES 6, 30C

Tree 6–20 m tall, trunk 10–20 cm in diameter, branchlets 

puberulous to tomentulose, glabrescent. Stipules caducous, 

not seen. Leaves compound, alternate, paripinnate; petioles 

(1–) 3–5(–9) mm long, puberulous; rachis (6.5–)9–14(–18) cm 

long, rachis wing narrow and erect without apical projections, 

contracted at each leaflet insertion, puberulous to tomentulose, 

rarely glabrous; leaflets (3–)6–9(–10.5) × (1–)2–3(–4) cm, (4–)6–

8(–12) jugate, sessile, elliptic to oblong–elliptic or obvate to 

oblanceolate–oblong, thinly coriaceous to coriaceous, the base 

inequilateral, the upper side shorter and attenuate, the lower 

rounded–obtuse, the apex acute or rounded to truncate and 

usually retuse to emarginate, usually puberulous on the salient 

midvein or at the blade base, more or less ciliolate along the margins, the lower surface micro-punctate. Inflorescences (5.5–)10–

20(–30) cm long, terminal racemes, elongating in fruit, the axis 

glabrous to pilosulose; pedicels (3–)5–9(–11) mm long, glabrous; 

bracts 23–30(–40) × 18–25 mm, oval to obovate, rounded apically, glabrous; bracteoles 15–30 × 8–15 mm, caducous, elliptic 

to obovate, united marginally in a tube enclosing the flower bud, 

acute apically, more or less carinate dorsally, joined marginally 

for 1⁄2–3⁄4 their length, glabrous. Flowers: hypanthium glabrous; 

stipe (2–)7–10 mm long; tube (11–)15–22 mm long and 5–7.5 

mm in diameter, cupular, green to red; sepals 4, (1.3–)1.8–2.6 × 

0.4–0.8(–1.1) cm, the adaxial one largest, narrowly elliptic to 

oblong, obtuse, red or greenish white, glabrous; petals 5, 12–20 × 

6–15 mm, the adaxial one ovate to triangular–ovate, the base 

auriculate to cordate, the apex acute, the other four petals 12–20 × 

6–9.5 mm, narrowly elliptic, acute to caudate-acuminate, usually bright red but sometimes white with red streaks, glabrous 

except apically ciliolate; stamens 3, red; filaments 4–7.5 cm long, 

more or less pilose; anthers 8–10.5 × 1.5–3 mm, linear to narrowly oblong, glabrous; staminodia 6, 10–20 mm long; tube 

0.8–1.5(–3) mm, basally united, except at the base of adaxial 

petal, more or less villose; gynoecium dark red with green margins, stigma capitate, style (1.2–)2–4(–6.8) cm long, glabrous 

except basally velutinous; ovary (5–)8–12(–15) × (1–)2–3 mm, 

narrowly elliptic to elliptic–oblong, velutinous, gynophore (1.5–)

3–7.5 mm long, adnate to upper surface of the hypanthium tube 

near its rim, glabrous except velutinous apically. Fruit 12–21 × 

4–6 cm, oblong–oblanceolate, bright red, velvety, transversely 

venose, a woody vertical flange parallel to legume valves along 

the placental suture, the flange 4–5 mm wide, ca. 6–10-seeded.

Distribution. Guyana; Suriname; Brazil: Amazonas, 

Roraima.

Phenology. Flowering and fruiting throughout the 

year (flowering specimens collected in all months except May 

and August, fruiting specimens lacking only for August and 

October).

Notes. This species is abundant in Guyana and is found 

in rainforest as well as savannah vegetation. On the 2004 expedition of author K.R., multiple individuals of the species were 

observed along the rivers and streams in the Rupununi region. 
1 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

On the return trip, the rainy season had begun and many of the 

trees were entirely submerged in water.

Additional specimens examined. BRAZIL. Amazonas: Territory of Roraima, Serra da Lua, Dormida, foothills of Serra 

da Lua, 2°25–29′N, 60°11–14′W, 15 Jan 1969 (fl., fr.), Prance et al. 

9287 (A, C, COL!, F!, G, INPA, K!, MG, MICH, MO!, NY!, P, R, 

S, U!, US!, VEN!); Roraima: Ilha de Maracá, Mun. Alto Alegre, 

3°24′N, 61°26′W, 14 Jul 1986 (fl.), Henderson & de Lima 604 

(K!, US!); Mun. Alto Alegre, SEMA Estação, 3°24′N, 61°26′W, 

14 Jun 1986 (fl.), Hopkins et al. 741 (K!); Mun. Boa Vista, 

Reserva Ecológica de Maracá, 14 Oct 1987 (fl., fr.), Lewis 1673

(K, 2 sheets!); SEMA Ecological Reserve, Ilha de Maracá, 3°20′N, 

61°33′W, 3 Jul 1988 (st.), Milliken & Bowles 389 (K!).

GUYANA. Cuyuni-Mazaruni: Essequibo River, 6–8 km 

downstream of Omai, 5°26′N, 58°42′W, 10–15 m, 1 Jun 1989 (fl., 

fr.), Gillespie 1579 (BRG!, K!, MO!, NY!, US!); Mazaruni River, 

Marshall Falls to several kilometers downstream, 6°20′0′′N, 

58°45′0′′W, 5 m, 26 Apr 1993 (fr.), Henkel 1969 (BRG!, K!, 

MO!, NY!, US!); Mazaruni River, small islands throughout 

river, 3.14 mi N of base camp, 6°8′51.2′′N, 58°53′59.7′′W, 31 m, 

13 Oct 2004 (fr.), Redden 3297 (BRG!, K!, MO!, NY!, US!); 

Mazaruni River, Marshall Falls, 6°16′54.4′′N, 58°42′58.4′′W, 

4 m, 17 Oct 2004 (fr.), Redden 3312 (BRG!, K!, NY!, US!). 

East Berbice-Corentyne: Along banks of Corentyne River, 

above Baba Grant sawmill, above Cow Falls, 5°00′N, 57°40′W, 

10–20 m, 17 Apr 1990 (fl., fr.), McDowell 2305 (BRG!, K!, 

MO!, NY!, US!); Corentyne River, between Cow Falls and Baba 

Grant sawmill, 5°00′N, 57°42′W, 10–25 m, 21 Apr 1990 (fl.), 

McDowell 2374 (BRG!, K!, MO!, NY!, US!); Corentyne River, 

Marabunta Creek, 4°56′N, 57°49′W, 10 m, 24 Mar 1995 (fl., fr.) 

Mutchnick 987 (BRG!, NY!, US!). Potaro-Siparuni: Iwokrama 

Forest Reserve, along Essequibo River between Turtle Mountain trail and Kurupukari Field Station, 4°41′N, 58°42′W, 

160 m, 18 Sep 1995 (fl.), Clarke 168 (BRG!, NY!, MO!, U!, 

US!); Essequibo River at Karupukari crossing, small island 0.25 

km SE of W bank landing, 4°40′N, 58°41′W, 55 m, 18 Apr 1992 

(fr.), Hoffman 1317 (BRG!, K!, MO!, NY!, US!); Iwokrama 

Forest Reserve, Karupukari Village, base camp 2 km N on Essequibo River, 04°41′N, 58°37′W, 60 m, 16 Mar 1997 (fr.), Mori 

24362 (BRG!, US!); Essequibo River, Kurupukari Falls, 4°40′N, 

58°40′W, 100 m, 7 Dec 1994 (fl.), Mutchnick 563 (BRG!, MO!, 

NY!, US!); Essequibo River, Kurupukari Falls and 2 km upstream, 

4°39′N, 58°39′W, 100 m, 12 Dec 1994 (fl.), Mutchnick 672

(BRG!, MO!, NY!, US!); Iwokrama Forest Reserve, Essequibo 

River, Akramukra Falls, 4°22′N, 58°30′W, 50 m, 16 Feb 1995 

(fl.), Mutchnick 758 (BRG!, K!, MO!, NY!, US!); Iwokrama 

Forest Reserve main field station, 4°40′17.7′′N, 58°41′7.1′′W, 

64 m, 27 Oct 2002 (fr.), Redden 1094 (BRG!, K!, NY!, U!, US!); 

Iwokrama Forest Reserve, Siparuni River heading toward the 

Burro-Burro River, between Pakatau Falls base camp and junction with Burro-Burro River, 4°44′57.8′′N, 58°57′14.2′′W, 56 m, 

1 Nov 2002 (fl. & fr.), Redden 1117 (BRG!, K!, US!); Iwokrama 

Forest Reserve main field station, on trial leading to Fairview 

Village, 4°40′17.7′′N, 58°41′7.1′′W, 64 m, 10 Nov 2002 (fl.), 

Redden 1179 (BRG!, K!, NY!, US!); 10 Nov 2002 (fl.), Redden 1180 (BRG!, K!, NY!, U!, US!); 10 Nov 2002 (fl.), Redden 

1181 (BRG!, K!, NY!, U!, US!). Upper Takutu–Upper Essequibo: 

Rupununi area, Karanambo, 3°53′0′′N, 59°35′0′′W, 110 m, 

17 Feb 1990 (fl.), Acevedo-Rodríguez 3266 (BRG!, K!, MO!, 

NY!, US!); Iwokrama Rainforest Reserve, Akromukru Transect 

at Akromukru Falls, Essequibo River, 4°29′N, 58°30′W, 70–90 m, 

18 Mar 1996 (f1.), Clarke 1327 (BRG!, K!, MO!, NY!, US!); 

Kuyuwini River, area near camp, 2°05′N, 59°17′W, 240 m, 

19 Jul 1996 (fl., fr.), Clarke 2314 (BRG!, K!, MO!, NY!, US!); 

Kwitaro River, landing at terminus of trail from Shea Village, 

2°54′N, 58°58′W, 135 m, 7 Feb 1997 (fl.) Clarke 3408 (BRG!, 

K!, NY!, MO!, U!, US!); Rewa River, 0–5 km S of confluence 

with Kwitaro River, 3°17′N, 58°45′W, 90 m, 25 Feb 1997 

(fl., fr.), Clarke 3916 (BRG!, K!, MO!, NY!, US!); Kuyuwini 

River, 0–5 km W of landing at terminus of trail from Karaudarnau and Parabara Savanna, 2°5′N, 59°14′W, 240 m, 31 May 

FIGURE 6. Geographic distribution of Paloue coccinea (M. R. 

Schomb. ex Benth.) Redden (dots) in Guyana and adjacent 

regions.
NUMBER 109 • 1 1

1997 (fr.), Clarke 5068 (BRG!, K!, MO!, NY!, US!); N. Rupununi, Apr 1968 (fr.), Davis 815 (K!); Iteballi Falls, Essequibo 

River, 1 Feb 1944 (fl.), Fanshawe F1667 (K!); South Rupununi 

savanna, Ikirap Creek along Marudi Road, 2°25′N, 59°15′W, 

250 m, 23 Dec 1993 (fl.), Henkel & James 3566 (BRG!, K!, 

MO!, NY!, U!, US!); South Rupununi savanna, Katambar Mtn. 

SE of Aishalton, 2°26′N, 59°08′W, 200 m, 17 Jan 1994 (fr.), 

Henkel 3662 (BRG!, K!, MO!, NY!, US!); Northern Rupununi savannas, Karanambo Ranch, trail SW of compound, 

3°45′12′′N, 59°19′36′′W, 90–120 m, 17 Feb 1992 (fl.), Hoffman 

979 (BRG!, K!, MO!, NY!, US!); Kuyuwini Landing, Kuyuwini 

River, 02°10′N, 59°15′W, 200 m, 1 Feb 1991 (fl., imm. fr.), 

Jansen-Jacobs et al. 2273 (K!, U!, US!); 2°05′N, 59°15′W, 150–

250 m, 12 Oct 1992 (fl., imm. fr.), Jansen-Jacobs et al. 2890

(BRG!, K!, U!, US!); Rupununi district, Kumukowau River, 

Camp 3, 2°56′N, 59°2′W, 160 m, 13 Feb 1994 (fl., fr.), JansenJacobs et al. 3731 (BRG!, K!, U!, US!); Ireng River base camp, 

4°08′38.9′′N, 59°38′28.3′′W, 136 m, 24 Mar 2004 (fl., imm. fr.), 

Redden 3018 (BRG!, K!, MO!, NY!, US!); Redden 3019 (BRG!, 

K!, MO!, NY!, US!); Ireng River near Ireng River base camp, 

4°08′25.8′′N, 59°37′57.9′′W, 113 m, 25 Mar 2004 (fl., imm. 

fr.), Redden 3031 (BRG!, K!, MO!, NY!, US!); Achimar Wau 

(Haimara creek), 2°54′21.1′′N, 59°16′14.6′′W, 155 m, 2 Apr 

2004 (fl., fr.), Redden 3045 (BRG!, K!, MO!, NY!, US!); Rupununi River, 2°49′5.5′′N, 59°31′41.6′′W, 113 m, 31 Mar 2004 

(fl., imm. fr.), Redden 3038 (BRG!, K!, MO!, NY!, US!); Rupununi savanna, 2°56′42.6′′N, 59°15′48.7′′W, 165 m, Redden 

3056, (fr.) Redden 3057 (BRG!, K!, MO!, NY!, US!); Kuyuwini 

River, 02°05′41.4′′N, 59°14′50.8′′W, 124 m, 14 Apr 2004 (fl., 

fr.), Redden 3082 (BRG!, K!, MO!, NY!, US!); Kuyuwini River, 

02°05′41.4′′N, 59°14′50.8′′W, 124 m, 14 Apr 2004 (fl.), Redden 3088 (BRG!, K!, MO!, NY!); Kuyuwini River, 2°5′41.4′′N, 

59°14′50.8′′W, 124 m, 14 Apr 2004 (fl., fr.), Redden 3089

(BRG!, K!, MO!, NY!, US!); Conservation International Concession on the Essequibo River, 3°53′58.1′′N, 58°22′50.2′′W, 97 

m, 19 Jan 2007 (fl.), Redden 5081 (BRG!, K!, MO!, NY!, US!); 

Basin of Kuyuwini River (Essequibo tributary), about 150 mi 

from mouth, 21–26 Nov 1937 (fl.), Smith 2521 (F!, K!, MAD, 

NY!, P, U!, US!); Pirara, 1841–1842 (fl.), Schomburgk 251 (K!, 

US!); Basin of Kuyuwini River (Essequibo tributary), about 150 

mi from mouth, 12 Feb 1938 (fl., fr.), Smith 3031 (K!, US!); 

Rupununi River, Sep 1948 (fl., fr.), Wilson Browne WB 136

(K!); Oronoque, New River, 25 Dec 1935 (fl.), Myers 3906 (K!); 

Kanuku foothills, 2 Feb 1952 (fl.), Guppy G172 (K!); without 

exact locality, without date (fl.), Appun 857 (K!).

SURINAME. Nickerie: Area of Kabalebo Dam project, 

4–5°N, 57°30′–58°W, 30–130 m, 6 Sep 1980 (fr.), Lindeman

et al. 172 (K!, US!). Sipaliwini: South side of Kuruni Island, riverbank E of trail from camp to canoe landing on S branch of 

Kuruni River around island, ca. 30 airline km E of confluence on 

Corantijn River, 3°22′12′′N, 57°20′36′′W, 180 m, 25 Nov 1994 

(fl., imm. fr.), Evans et al. 1998 (MO!, U!, US!); Wonotobo, Corantyn, 13 Oct 1916 (fr.), Stahel & Goggryp 2861 (U!).

4. Paloue duckei (Huber) Redden comb. nov.

Elizabetha duckei Huber, Bol. Mus Goeldi 7: 291. 1913. Type. Colombia. 

Amazonas: Puerto Cordoba, Rio Japura (Caqueta) near the Cupati 

Rapids, 14 Nov 1912 (fl.), A. Ducke 12245 (holotype: MG; isotypes: 

BM!, G, P!, RB, US!).

FIGURES 7, 8

Small tree to 10 m tall, branchlets pilose or pilosulose. 

Stipules ca. 6 cm long, caducous, thin, oblanceolate, the apex 

reflexed, puberulous externally and ciliolate on margins. 

Leaves compound, alternate, paripinnate, lanceolate; petioles 

1.5–3.5 mm long, terete, pilosulose, rachis 9–14.5 cm long, 

rachis wing narrow, apical projections at each leaflet insertion, 

glabrous; leaflets 17–25 × 3–5 mm, 18–30-jugate, sessile, lanceolate to narrowly oblong, base inequilaterally truncate, the 

apex rounded, slightly retuse, micro-mucronulate, the mucro 

caducous, micro-punctate beneath, glabrous; venation subobscure. Inflorescences 5–7 cm long, open, axillary, sessile 

racemes, the axis puberulous but soon glabrescent; bracts ca. 

6 × 4 mm, caducous, obovate, glabrous; bracteoles 10–11.5 × 

4–4.5 mm, caducous, acute, slightly carinate dorsally, united 

unequally, the union more complete on the adaxial side of the 

flower, red, glabrous; pedicels 3–6 mm long, often puberulous 

but glabrescent. Flowers: hypanthium 13–17 mm long, 5.5–

8.5 mm wide, usually glabrous but occasionally a pilose patch 

present on tube; stipe 3–9 mm long; tube (12–)20–21 mm long, 

4–7 mm in diameter at the apex, gradually tapering to the 

stipe, red; sepals 4, 1.3–1.7 × 0.5–0.9 cm, oblong, glabrous; 

petals 5, (12–)14–17 × (3.5–) 5–8.5 mm, about equal in size 

and shape, elliptic, glabrous except ciliate apically; stamens 3, 

red, “very long erect” (fide Ducke, 1934), filaments 3.7–6.3 cm 

long, sparingly villose; anthers 5.5–8.5 × 2.5–3 mm, more or 

less elliptic; staminodia 6, 0.5–1 mm long, highly reduced; tube 

united for 0.5–1 mm; gynoecium glabrous; glabrous; ovary 

8–10 × 1.5–2.5 mm, narrowly oblong; gynophore 2–4 mm 

long, style 3–4.5 cm long, stigma capitellate. Immature fruit

12–13 × 3–3.5 cm, flat, oblong, narrowly alate, glabrous, 

4–8-seeded.

Distribution. Brazil: Amazonas, Belém; Colombia: 

Amazonas.

Phenology. Collected in flower in March, August, and 

November, and in fruit in August and September.

Additional specimens examined. BRAZIL: Amazonas: Igarape Jacitari, 01°23.197′N, 68°10.377′W, 19 Aug 

2008 (fl., fr.), Acevedo-Rodríguez 14648 (K!, US!); Belém, in 

horto Musei e Rio Caquetá, Puerto Córdoba, anno 1912 introducta, Ducke 767 (F, K!, MO!, NY!, R, US!); culta in Horto 

Musei Paraensis (e regione cataractarum inferiorum fluminis 

Caquetá, Colombia), 4 Mar 1926 (fl.), Ducke s.n. (RB-20311, 

K!, NY!, P, R, U!, US!); Rio Icana, Tunui, 28 Mar 1952 (fl.), 

Fróes 28100 (NY!, US!). Belém: Jardim Botanico do Museu 

Goeldi, 3 Sep 1942 (fr.), Archer 7626 (US!).
1 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 7. Paloue duckei (Huber) Redden: A, flowering branch; B, detail of upper surface of leaflets; C, detail 

of lower surface of leaflets; D, glands on lower surface of leaflets; E, flower. Scale bar length for each: A = 3 cm; 

B, C = 7 mm; D = 1.6 mm; E = 1.5 cm. Drawn from Ducke 20311 (K) by Lucy Smith. Copyright © The Board of 

Trustees of the Royal Botanic Gardens, Kew.
NUMBER 109 • 1 3

5. Paloue durissima (Ducke) Redden comb. nov.

Elizabetha durissima Ducke, Trop. Woods 37: 26. 1 Mar 1934. Type. Brazil. Amazonas: Parintins, Lago Jose-Assu, 16 Sep 1932 (fl.), A. Ducke 

23729 (holotype: RB; isotypes: F!, G, IAN, K!, MAD, NY!, P!, R, U!, 

US!).

FIGURES 9, 10, 29A

Tree ca. 25 m tall, the branchlets pale brown to black, pilosulose. Stipules ca. 25 × 5 mm, caducous, membranous, acute, 

narrowly oblanceolate, strigulose externally, ciliolate on the 

apical half. Leaves compound, alternate, paripinnate, oblong to 

lanceolate; petiole 1.5–2 mm long, pilosulose; rachis 7.5–11.5 cm 

long, rachis wing narrow, puberulous with apical projections 

at each leaflet pair insertion, the axis pilosulose; leaflets 1.6–

1.8 × 0.2–0.3 mm, 34–41-jugate, sessile, narrowly lanceolate–

oblong, slightly arcuate, the base inequilaterally obtuse, the 

apex acute, ciliolate, the venation sub-obscure. Inflorescences

2–4 cm long, short axillary racemes, the axis densely tomentose; pedicels 2–3 mm long; bracts 7–9 × 5–10 mm, caducous, 

rounded, obtuse, white, strigulose externally and ciliolate on 

margins; bracteoles 5–8.5 mm long, united almost to their 

apex, thin, acute, externally sparingly tomentulose, pilosulose, 

or tomentose. Flowers: hypanthium, pilosulose to tomentulose; 

stipe 3.5 mm long; tube ca. 4 mm long and 4.5 mm in diameter, cupular, white; sepals 8–10 × 3–6 mm, narrowly ovate to 

lanceolate, white, glabrous except ciliolate near apex; petals ca. 

8 × 6.5 mm, the dorsal one triangular-ovate, acute, the other 

four narrower, sublinear-oblong, 9–9.5 × 1.5–2 mm, acuminate, white, glabrous; stamens 3, white; filaments 2.3–2.5 cm 

long, sparingly villose; anthers ca. 3 × 1.5 mm, elliptic, glabrous; staminodia 6; gynoecium red; ovary ca. 5 × 1.5 mm, 

oblong–elliptic, glabrous; gynophore 2 mm long, style 1.8–2.5 

cm long, stigma capitate. Fruits 10–15 × 4–4.5 cm, glabrous, 

ca. 3–6-seeded.

Distribution. Guyana; Brazil: Amazonas and Pará.

Phenology. Collected in flower in September, November, and January, and in fruit in April.

Note. A specimen was collected by Ducke (81000) in 

the Rio de Janeiro Botanical Garden in October 1952 (Cowan, 

1976).

Additional specimens examined. GUYANA. 

Upper Demerara-Berbice: Mabura Hill, 30 km S on Mabura 

Hill-Kurupukari Road, 5°05′N, 58°44′W, 75–85 m, 2 Jan 1996 

(fl.), Clarke 753 (BRG!, MO!, NY!, US!). Potaro-Siparuni:

Iwokrama Forest Reserve, base of Turtle Mountain in gap clearing, 4°43′56.2′′N, 58°44′1.6′′W, 20–25 m, 9 Nov 2002 (fl.), 

Redden 1171 (BRG!, K!, MO!, NY!, US!), Redden 1178 (BRG!, 

MO!, NY!, US!); Upper Takutu, just below Pakatau Falls, 

4°45′10.00′′N, 59°01′25′′W, 26 Apr 1992 (fr.), Pennington et al. 

400 (K!).

BRAZIL: Pará: Estacão Ecologia do Jari, Projeta Reserva 

Genetica, SEMA, 00°75′S, 52°30′W, 17 Oct 1997 (fl.), Beck

et al. 139 (K!, US!).

6. Paloue emarginata (R. S. Cowan) Redden comb. nov.

Paloveopsis emarginata R. S. Cowan, Brittonia 8: 4. 1957. Type. Guyana. 

Side of Bunawau Creek, 31 Oct 1952 (fl.). N. Guppy 513 (holotype: 

NY!; isotypes: F!, K!, US!).

FIGURES 11, 29D

Medium-sized tree 11 m tall, 20 cm in diameter, branchlets and petiole pilose, later becoming glabrous. Stipules 9 mm 

long, caducous, glabrous. Leaves simple (unifoliolate), alternate; 

petiole 3–4 mm long, glabrous, leaf blade 5.5–7.5 × 2.5–3.5 cm, 

elliptic, base acute, apex acute and emarginate, pubescent on the 

midvein on upper surface of leaf blade, upper surface glabrous 

and minutely punctuate, midvein salient, venules conspicuous. 

Inflorescences axillary or terminal racemes, 3–5 cm long, bracts 

2 × 1.5 mm, caducous, sessile, glabrous, minutely ciliolate on 

margins; bracteoles connate, 5 × 2.5 mm, obovate, apex ciliolate 

on margins; pedicels 5.5–6 mm long, glabrous. Flowers: hypanthium glabrous, 5 mm long, 4 mm in diameter, funnel-shaped, 

FIGURE 8. Geographic distribution of Paloue duckei (Huber) 

Redden (dots) in Brazil.
1 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 9. Paloue durissima (Ducke) Redden: A, flowering branch; B, upper portion of leaf; C, detail of upper 

surface of leaflets; D, detail of lower surface of leaflets; E, glands on lower surface of leaflets; F, bud; G, bud with 

calyx and corolla removed to show stamen packaging; H, gynoecium showing stipe attachment to hypanthium 

wall; I, petals; J, sepals. Scale bar length for each: A = 3 cm; B, F–J = 7 mm; C, D = 2.5 mm; E = 1 mm. Drawn 

from Redden 1171 (K) by Lucy Smith. Copyright © The Board of Trustees of the Royal Botanic Gardens, Kew.
NUMBER 109 • 1 5

7. Paloue fanshawei (R. S. Cowan) Redden comb. nov.

Elizabetha fanshawei R. S. Cowan. Proc. Kon. Nederl. Akad. Wetensch. Ser. 

C. 79: 323–334. 1976. Type. Guyana. 107 mi along Bartica–Potaro 

Road, 16 Nov 1943 (fl., fr.), D. B. Fanshawe 1498 (holotype: NY!, 

image at K!, isotype K!).

FIGURES 12, 13

Tree 12–15 m tall, trunk 15–20 cm in diameter, branchlets puberulous. Stipules caducous, not seen. Leaves compound, 

alternate, paripinnate; petioles 3–4 mm long, minutely pubescent, rachis 5–6.5 cm long, rachis wing narrow and erect without 

apical projections at each leaflet insertion, minutely pubescent; 

leaflets 4–6-jugate, sessile, the blade oblong to elliptic–oblong, 

base inequilateral, the upper acute, lower rounded-obtuse, apex 

rounded, obtuse and retuse, glabrous above, below micropunctate and near to the base a little tuft of long pilose hairs along 

the salient midvein. Inflorescences axillary racemes, 8 cm long, 

glabrous; bracts caducous, not seen; bracteoles oval, 14 × 9 mm, 

united marginally in a tube enclosing the flower bud, glabrous; 

stipe 4.5–5 mm long, glabrous; sepals 10 × 3.5–5.5 mm, glabrous, elliptic, imbricate; petals 5, equal, 6.5–8 × 1–1.5 mm, 

linear-lanceolate, apex acute, punctate, long-ciliate along margins; stamens 9, 3 fertile, filaments of the fertile stamens ca. 

21 mm long, villose, anthers oblong, 4.5 × 1 mm, staminodes 

3–4.5 mm long, tapering toward tip, villose, filaments of stamens and staminodes connate at base for 1–2 mm; ovary 4 × 1.5 

mm, glabrous, oblong, stipe 1.5 mm long, glabrous; style 16 mm 

long, glabrous; stigma capitate. Fruit a legume, (11–)13–18 × 

4.5–5.5 cm, oblong but somewhat wider toward the apex, ca. 

6–8-seeded.

Distribution. Guyana; Suriname; Brazil: Amazonas.

Phenology. Collected in flower in June and October 

and in fruit in June and August.

Additional specimens examined. BRAZIL. Amazonas: Vicinity of Rio Uatumã, Rio Pitinga, margem direita do 

rio, 29 Aug 1979 (fr.), Cid et al. 916 (K!, NY!, US!).

SURINAME. Sipaliwini: Sipaliwini River, 0–1 km downstream from Kwamalasamutu Village, 2°22′31′′N, 56o46′31′′W, 

213 m, 17 Jun 2004 (fl., fr.), Hoffman 5924 (BBS, U!, US!).

FIGURE 10. Geographic distribution of Paloue durissima

(Ducke) Redden (dots) in Guyana and Brazil.

FIGURE 11. Geographic distribution of Paloue emarginata (R. S. 

Cowan) Redden (dots) in Guyana, Suriname, and Brazil.
1 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 12. Paloue fanshawei (R. S. Cowan) Redden: A, flowering branch; B, upper surface of leaflet; 

C, lower surface of leaflet; D, glands on lower surface of leaflet; E, fruiting branch. Scale bar length for each: 

A, E = 3 cm; B, C = 1.5 cm; D = 1 mm. A–D drawn from Hoffman 1069 (K); E drawn from Fanshawe 1044 

(K) by Lucy Smith. Copyright © The Board of Trustees of the Royal Botanic Gardens, Kew.
NUMBER 109 • 1 7

FIGURE 13. Geographic distribution of Paloue fanshawei (R. S. 

Cowan) Redden (dots) in Guyana.

pedicel 14–16 mm long. Flowers: hypanthium glabrous, stipe ca. 

8 mm long, tube ca. 6.5 mm long; sepals green, ca. 15 × 6–8 mm; 

petals 5, ovate, adaxial petal largest, ca. 27 × 16 mm, white, villose at base, the reduced petals linear, ca. 2 mm long; stamens 

9, filaments 5.5 cm long, fused into a tube 3.5 mm long except 

at adaxial petal, villose toward base; ovary velutinous, oblong, 

ca. 6 × 2 mm, stipe 3 mm long, velutinous; style pilose at base. 

Fruit a legume, pilose to minutely pubescent, flat, ca. 5.5 cm 

long, 4–6-seeded.

Distribution. Endemic to Guyana.

Phenology. Collected in flower in November, February and March, and in fruit in January, February, May, and 

October.

Additional specimens examined. GUYANA.

Mahdia River, Potaro River, 107 mi Bartica–Potaro Road, 

18 Jan 1943 (fr.) Fanshawe F1044 (K!); Cuyuni-Mazaruni:

S. Pakaraima Mts., at “Corona Falls,” ca. 20 km NW of Karasabai Village, 4°09′N, 59°41′W, 150 m, 27 Feb 1992 (fl.), Hoffman 1069 (BRG!, K!, MO!, NY!, U!, US!); Pakaraima Mts., 

base camp 8.6 km NE of Imbaimadai on Partang River tributary, 

5°46′N, 60°15′W, 650 m, 23 May 1992 (fr.), Hoffman 1814

(BRG!, MO!, NY!, US!); base camp, small tributary of Partang 

River, 11.4 km NE of Imbaimadai, 5°48′N, 60°15′W, 650 m, 

26 May 1992 (imm. fr.), Hoffman 1897 (BRG!, K!, MO!, NY!, 

US!); West branch of Eping River, 6°00′0′′N, 60°10′0′′W, 122–

152 m, 7 Feb 1991 (fr.), McDowell 3895 (BRG!, K!, MO!, NY!, 

US!); Pakaraima Mts., Mazaruni River, 9.21 mi W of Imbaimadai to Kamarang, near Jawalla and Baimapai, 5°43′51.1′′N, 

60°35′15′′W, 498 m, 5 Feb 2004 (fr.), Redden 1722 (BRG!, K!, 

MO!, NY!, US!); Pakaraima Mts., Mazaruni River between 

Imbaimadai and Kamarang, near Jawalla and Baimapai; 20.2 

GPS mi W and downstream of Imbaimadai, 5°43′51.1′′N, 

60°35′15′′W, 498 m, 6 Feb 2004 (fr.) Redden 1733 (BRG!, 

K!, MO!, NY!, US!); Mazaruni River base of Arawai Falls, 

6°20′41.2′′N, 60°34′29.1′′W, 133 m, 9 Oct 2006 (fr.), Redden

et al. 4648 (K!); Warashema Mountain Range, 6°30′41.7′′N, 

60°25′3.7′′W, 509 m, 2 Mar 2007 (f1.), Redden 5429 (BRG!, 

K!, MO!, NY!, US!).

8. Paloue × grahamiae (R. S. Cowan) Redden, comb. et stat. nov.

(P. induta × P. coccinea)

Elizabetha grahamiae R. S. Cowan, Proc. Kon. Nederl. Akad. Wetensch. 79: 

323–334. 1976. Type. Guyana. Mazaruni River, Marshall Falls, 28 Apr 

1960 (fl.), V. Graham 355 (holotype: K!).

FIGURES 14, 15, 30D

Tree 10 m tall, branchlets pubescent but becoming glabrous. 

Stipules caducous, not seen. Leaves compound, alternate, imparipinnate, with 1–3 lateral leaflet pairs, and a single terminal leaflet; petiole ca. 4.5 mm long, pubescent, rachis ca. 10 mm long, 

narrowly winged, without apical projection at leaflet insertion, 

the erect wing minutely pubescent; leaflets coriaceous, elliptic–

lanceolate, sessile, glabrous, large terminal leaflet 11–12.5 × 4–5 

cm, apex acute, base equal, obtuse, with paired crater-like glands 

at the base of the lamina, lateral leaflets 2.5–4 × 1.5–2 cm, base 

inequilateral, acute, apex acute, midvein salient on lower surface, secondary venation slightly raised on lower surface. Inflorescences terminal or axillary racemes, 6.5–9 cm long, glabrous; 

bracts ca. 14 × 10 mm, cucullate, oval, glabrous, apex rounded; 

bracteoles ca. 15 mm long, obtuse to acute; pedicels glabrous, 

11 mm long. Flowers: hypanthium glabrous, greenish, stipe 

10 mm long, tube ca. 15 × 7 mm; sepals reflexed, narrowly elliptic to oval, glabrous, 17–18 × 8–11 mm, reddish; petals 5, red, 

dorsal petal ca. 12 × 10 mm, elliptic, ciliolate at apex, 4 other 

petals, long-acuminate, 11–15 × ca. 1.5 mm, apex ciliate; stamens 9, glabrous, filaments ca. 6.5 cm long, fused tube 3 mm 

long, anthers of three different sizes; ovary ca. 10 × 2.5 mm, velutinous, narrowly elliptic, stipe ca. 4 mm long, slightly pubescent; 

style glabrous except at base. Fruit a legume, 12–21 × 4–6 cm, 

pubescent, ca. 4–8-seeded.

Distribution. Endemic to Guyana, this hybrid is 

found where P. coccinea co-occurs with P. induta or P. riparia.
FIGURE 14. Field drawing of Paloue × grahamiae (R. S. Cowan) Redden by Violet Graham. Illustration reproduced courtesy of the Royal Botanic Gardens, Kew.
NUMBER 109 • 1 9

Essequibo: Kuyuwini River, 2°05′41.4′′N, 59°14′50.8′′W, 124 m, 

14 Apr 2004 (fl., fr.), Redden 3090 (BRG!, K!, MO!, NY!, U!, 

US!); Kuyuwini River, 2°04′19.1′′N, 59°10′57.9′′W, 243 m, 

14 Apr 2004 (fl.), Redden 3104 (BRG!, K!, MO!, NY!, U!, US!).

9. Paloue guianensis Aubl., Hist. Pl. Guiane 1: 365–367, pl. 141. 

1775. Lectotype (here designated). French Guiana: “Forest 

of Guiane near the town known by the name Saint-Régis,” 

J. B. C. F. Aublet s.n. (BM!).

FIGURES 16, 17, 29E

Small shrub or tree to 10 m tall, branchlets gray/brown, glabrescent. Stipules caducous, linear to strap-like. Leaves simple, 

alternate, entire, blade lanceolate–oblong, 10–16 × 2.5–6 cm, 

apex acuminate, base acute to rounded to auriculate, usually 

with paired crater-like glands at the base of the lamina but 

sometimes these not present, blade coriaceous, glabrous; petioles 3–5 mm long. Inflorescences terminal or axillary racemes, 

11–14 cm long; bracts orbiculate, 1 cm long, persistent or caducous; bracteoles connate, 6–10 cm long, bilobed, rounded, persistent; pedicels 6–8 mm long. Flowers: hypanthium 1.5–2 cm 

long including the stipe; sepals deep red, 15–20 × 7 mm, 

rounded at the apex, reflexed; petals 5, red, the three superior 

petals nearly equal, spatulate, 20–30 × 7 mm, or the two lateral 

petals much smaller, the two inferior petals filiform, 2–3 mm 

long, margins glabrous; stamens 9, red, filaments 4–6 cm long, 

connate equally to form a tube, sparsely long-pilose, anthers 

linear-oblong, glabrous to sparsely pilose, 3–5 mm long; ovary 

1 cm long, glabrous to tomentose, stipitate, style 3–6 cm long, 

stigma capitate. Fruit a legume, 20–23 × 3–5 cm, oblong, stipitate, transversely veined, acuminate, red-brown tomentose, ca. 

4–8-seeded.

Distribution. Guyana; French Guiana; Suriname; Brazil: Amapa, Pará.

Phenology. Flowering from October to April, fruiting 

throughout the year (fruiting collections only unknown from 

January, February, and May).

Notes. Specimens were not cited in Aublet’s protologue 

of Paloue guianensis, but a detailed drawing was included. After 

Aublet’s death in 1778, his herbarium was divided and put up for 

sale, with the main part now housed in the Banks Herbarium of 

the Natural History Museum (BM), and in Paris (P), with other 

specimens at BR, F, LIV, MO, UPS, and W (Stafleu and Cowan, 

1976). One specimen at BM was assumed to be Aublet’s type of 

Paloue guianensis and is labeled as such, but no documentation 

as to who designated it as the type is included. Pulle (1906) and 

Sandwith (1937) both commented on this specimen, and Sandwith recognized the similarity between the Aublet specimen at 

BM and Aublet’s drawing, stating “The specimen which may 

with justification be taken as Aublet’s type shows a flower with 

three long, subequal petals which accords with his figure.” No 

formal designation of this specimen as the lectotype has been 

published, so we have made that designation here.

FIGURE 15. Geographic distribution of Paloue × grahamiae

(R. S. Cowan) Redden (dots) in Guyana.

Phenology. Collected in flower in April and November, and in fruit in April.

Notes. Phylogenetic analyses based on morphology and 

DNA sequence data from the plastid trnL intron and nuclear 

ITS provided evidence that P. grahamiae may be of hybrid origin 

(Redden, 2008; Redden et al., 2010). To investigate this further, 

cloning experiments were conducted to check for multiple copies of ITS in the putative hybrid. Cloning yielded two distinct 

copies of ITS from the hybrid, which sorted with the putative 

parents, P. induta and P. coccinea, suggesting a recent origin of 

the hybrid. A specimen of P. × grahamiae collected in 2007 was 

growing in close proximity to individuals identified as P. induta

and P. coccinea.

Additional specimens examined. GUYANA.

Cuyuni-Mazaruni: Mazaruni River, Marshall Falls, ca. 20 mi 

from Bartica, 6°17′N, 58°43′W, 18 Nov 2005 (fl.) McDowell 

5744 (BRG!, US!); 6°15′47.2′′N, 58°45′1.6′′W, 6 m, Redden 

3307 (BRG!, K!, MO!, NY!, U!, US!). Upper Takutu-Upper
2 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 16. Paloue guianensis Aubl., reproduced from Aublet (1775: atlas 1, pl. 141): 1, inflorescence; 2, bracteoles; 

3, hypanthium; 4, petal; 5, stamen; 6, ovary; 7, fruit.
NUMBER 109 • 2 1

Additional specimens examined. BRAZIL. 

Amapa: Rio Oiapoque, Cachoeira Grande Roche, 3°48′N, 

51°53′W, 17 Jul 1960 (fr.), Maguire et al. 47045 (U!); Rio 

Oiapoque, ilha perto da cachoeira do Carratá, 1 Feb 1950 (fr.), 

Fróes 25762 (K!); Pará: Jacundá, 3 km toward Jatobal; area to 

be flooded by Tucuruí Dam, 23 Oct 1977 (fl.), Silva et al. AS152

(K!, MO!, U, US!).

GUYANA. Upper Demerara-Berbice: Mabura Hill area, 

Camudi compartment, 5°20′N, 58°10′W, 15 Mar 1996 (fl., fr.), 

Roberts 2 (U!).

SURINAME. 1862 (imm. fl.) Kappler 81 (L!). Marowijne: 

Ad ripas fluv. Marowijne, 14 Feb 1949 (fl.), Lanjouw & Lindeman 2034 (K!, U!); Nickerie: Area of Kabalebo Dam project, 

30–130 m, 20 Nov 1981 (fl.), Lindeman & Roon 874 (BBS, K!, 

U!); Sipaliwini: Gonini, 20 Feb 1918 (fr.), Gonggryp 3737 (U!).

FRENCH GUIANA. Cayenne: Sinnamary: Saut L’AutelBassin du Sinnamary, 4°45′N, 53°6′W, 25 m, 24 Mar 1994 (fl.), 

Bordenave 871 (NY!, P!, U!, US!); D. Z. de Saut-Dalles-Bassin du 

Sinnamary, 4°33′N, 52°53′W, 60 m, 7 Jan 1992 (fl.), Hoff 7211

(CAY!, NY!, P!, U!, US!); Cayenne (fl.), von Rohr s.n. (BM!); 

Crique Gabaret-Bassin de l’Oyapock, 3°55′N, 51°48′W, 15 m, 

11 Apr 1988 (fl., fr.), Cremers 9864 (CAY!, NY!, P, U!, US!); Trésor Reserve, 4°35′N, 52°16′W, no date (fl.), Ek 1579 (U!); Regina 

Region, eastern plateau of Montagne Tortue, 11 km WNW of 

Approuague River, 4°18′N, 52°22′W, 200–450 m, 14 Jun 1988 

(fr.), Feuillet 10065 (U!); zone Crique Probert, entre Saint-Georges 

et le pont transfrontalier sur l’Oyapock, 3°52′50′′N, 51°48′46′′W, 

6 m, 20 Dec 2005 (fl.), Granville & Crozier 17222 (CAY, K!, US); 

St. Georges de l’Oyapock, Crique Gabaret; en amont et en aval 

du Saut Emerillon, 27 Dec 1978 (jv. fr.), Grenand 1781 (US!); Rio 

Oiapoque, 3°43′N, 51°55′W in French Guiana, opposite Colonia Agricola do Oiapoque, Brazil, about 4 km N of Cricu River, 

13 Aug 1960 (fr.), Irwin & Westra 47476 (K!, NY); Maroni, without date (fl.), Mélinon 18 (K!); bords de la Rivière du Maroni, 

1862 (fl.), Mélinon s.n. (K!); Mariaflor, Oyapock, 2 Dec 1965 (fl.), 

Oldeman 1835 (U!); Riviére Comté, sur la creique Bagot, branche 

Nord à environ 3,700 km en amont de son embranchement, 

18 Dec 1968 (st.), Oldeman B-2054 (U!); Riviére Camopi (affluent de l’Oyapock), rives du Saut Ouayeouarou, 9 Feb 1968 (fl.), 

Oldeman & Sastre 115 (U!); Crique Alikene about 0–50 km from 

its confluence with River Camopi, ca. 3°10–20′N, 52°28–32′W, 

30 Sep 1960 (fl., fr.), Pires 48561 (K!, U!); Fleuve Approuague, 

riviere Arataye, Saute Pararé, 29 Oct 1978 (fl., fr.), Poncy 183

(P!, U!, US!); Régina, Réserve Naturelle Des Nouragues, Camp 

Arataï, Rives de la riviére Arataye, 3°59′30′′N, 52°35′46′′W, 

100 m, 20 Feb 2003 (fl., fr.), Poncy 1666 (K!, P!, US!); Crique 

Beiman sur le Maroni, 4°37′N, 54°20′W, 22 Feb 1984 (jv. fr.), 

Prévost 1717 (US!); Crique Gabrielle, 4°43′32′′N, 52°17′59.2′′W, 

57 m, 15 Apr 2007 (fr.), Redden 5968 (CAY!, US!); Fleuve 

Moroni, Pompidou-Papaichton, 27 Dec 1978 (fl.) Sastre 6470

(U!); Antécume Pata, 3°15′N, 54°10′W, 134 m, 21 Feb 1991 

(fl.), Veth & Manu 32 (U). Roura: On road at far side of Cacao 

leading to town water reservoir, ca. 200–500 m from Cacao, 

4°33′37.1′′N, 52°27′46.8′′W, 66 m, 1 Nov 2009 (fl.), Mori et al. 

27358 (CAY!, NY!, US!).

10. Paloue induta Sandwith, Bull. Misc. Inform. Kew 1937(2): 

106–107. 1937. Type. Guyana (as British Guiana). Upper 

Essequibo River: Mataruki River, between camps 1 and 2, 

3 Dec 1935 (fl.), J.G. Myers 5801 (holotype: K!).

10a. Paloue induta subsp. induta

FIGURES 18, 30A

Medium-sized tree, 25–35 m tall, branchlets cinnamoncolored, minutely pubescent to glabrous. Stipules caducous, not 

seen. Leaves simple, alternate, entire, up to 11 × 3–5 cm, blade 

oblong–lanceolate or lanceolate, apex acuminate, base slightly 

rounded to cuneate with a pair of crater-like glands at the base of 

the lamina, one either side of the main vein, coriaceous, glabrous, 

petioles 3–4 mm long. Inflorescences solitary in axils or terminal 

racemes, 4–6 cm long, rachis and pedicels with short, spreading, somewhat glossy, golden hairs; bracts widely ovate, apex 

round-obtuse, ca. 8 mm long, outer surface densely pubescent, 

FIGURE 17. Geographic distribution of Paloue guianensis Aubl. 

(dots) across the Guiana Shield.
2 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

caducous; bracteoles connate, bilobed, rounded, persistent, 

7–10.5 mm long, with a dense, pearl-gray pubescence; pedicels 

6–9 mm long. Flowers: hypanthium 12–25 mm long including 

stipe, densely pubescent with ascending, curved, short trichomes; 

sepals deep red on inner surface, green on outer surface, 1.3–1.5 × 

0.6–1.3 cm, broadly ovate or oblong, obtuse, reflexed; petals red, 

glabrous, oblong or oblong-linear, largest petal acute, 13–14 × 

3–5 mm, two medium-sized petals with obtuse apex, 3.5–5 × 

ca. 1.25 mm, two smallest petals with obtuse apex, about 1.8 × 

0.75 mm, margins glabrous; stamens 9, red, filaments 4.8–5.3 

cm long, uniformly connate, covered with long trichomes that 

weakly flex upward, anthers glabrous, 7–7.5 × 1.5 mm, the apex 

terminating in a little mucro that is round to obtuse, about 0.2 

mm long; gynoecium tomentose, densely covered in yellow-white 

trichomes, stipe glabrous, style 5–6.5 cm long, base covered with 

trichomes similar to those on ovary. Fruit a legume, 11–15 × 2–4 

cm; ca. 4–8-seeded.

Distribution. Guyana; Suriname.

Phenology. Flowering from September to December 

and in April, fruiting from March to May.

Notes. This taxon differs from all the other species of 

Paloue in that the entire inflorescence is covered with trichomes 

and the ovary is distinctly villous to tomentose. P. induta subsp. 

induta is morphologically similar to P. riparia Pulle and P. guianensis Aubl., but P. riparia has smaller flowers and bracteoles 

and its ovaries are glabrous, and P. guianensis has a distinctly 

different petal configuration.

Additional specimens examined. GUYANA.

Upper Takutu–Upper Essequibo: Kamoa River, 0–6 km W of 

camp, 1°31′50′′N, 58°49′40′′W, 9 Nov 1996 (fl.), Clarke 3010

(BRG!, K!, U!, US!); Kamoa Mts., 2 km S of Kamoa River, 

1°31′4′′N, 58°48′54′′W, 13 Nov 1996 (fl.), Clarke 3207 (BRG!, 

K!, NY!, US!); Essequibo River, between Camp (at confluence 

of Essequibo River and Onoro Creek) and Konashen Rapids, 

1°35′N, 58°37′W, 18 Nov 1996 (fl.), Clarke 3289 (BRG!, NY!, 

U!, US!); Sipu River, area near camp, 1°25′N, 58°57′W, 240 m, 

11 Sep 1998 (fl.), Clarke 7687 (BRG!, NY!, US!); Sipu River, 0–3 

km W of camp, 1°25′17′′N, 58°57′7′′W, 240 m, 12 Sep 1998 (fl.), 

Clarke 7790 (BRG, NY, US); Acarai Mts., Sipu River 8–10 km 

from juncture with Essequibo River, 0°25′N, 58°51′W, 250 m, 

12 Mar 1994 (fr.) Henkel et al. 5108 (BRG!, K!, MO!, NY!, 

U!, US!); Kuyuwini River, 2°5′41.4′′N, 59°14′50.8′′W, 124 m, 

14 Apr 2004 (fr.), Redden 3092 (BRG!, K!, NY!, US!); Parabara 

Village, 1.21 mi SW of village, 2°5′13.4′′N, 59°15′22.6′′W, 266 

m, 16 Apr 2004 (fl., fr.), Redden 3099 (BRG!, K!, NY!, US!); 

Parabara Village, Parabara landing, 2°5′54.5′′N, 59°14′58.2′′W, 

238 m, 16 Apr 2004 (fl.), Redden 3103 (BRG!, K!, NY!, US!); 

Essequibo River, Gunn’s region, 1°33′55.7′′N, 58°38′54.7′′W, 

236 m, 30 Apr 2004 (fr.), Redden 3154 (BRG!, K!, NY!, US!); 

Essequibo River, Gunn’s region, 1°34′6.6′′N, 58°39′25.6′′W, 235 

m, 30 Apr 2004 (fr.), Redden 3155 (BRG!, K!, NY!, US!); Sipu 

River, 1°25′28.2′′N, 58°51′40.5′′W, 225 m, 3 May 2004 (fr.), 

Redden 3160 (BRG!, K!, NY!,US!); Sipu River, 1°25′28.2′′N, 

58°51′40.5′′W, 225 m, 3 May 2004 (fr.), Redden 3169 (BRG!, 

K!, NY!,US!); Chodikar River, 1°24′51.5′′N, 58°47′44.5′′W, 242 

m, 4 May 2004 (fr.), Redden 3189 (BRG!, K!, NY!, US!); Basin 

of Essequibo River, near mouth of Onoro Creek, 1°35′N, 15–24 

Dec 1937 (fl.), Smith 2652 (F!, K!, MO!, NY!, US!). PotaroSiparuni: Micobe, along bank of Tiger Creek, 5°20′N, 59°30′W, 

200 m, 20 Oct 1991 (fl.) Tiwari 488 (NY!) Micobe, 5°20′N, 

59°30′W, 200 m, 21 Oct 1991 (fl.), Tiwari 511 (NY!, US!); 

Micobe, along bank of Tiger Creek, 5°20′N, 59°30′W, 200 m, 

21 Oct 1991, (fr.) Tiwari 517 (US!)

SURINAME. Sipaliwini: Gonini River, Sep 1903 (fl., imm. 

fr.), Versteeg 248 (U!).

10b. Paloue induta subsp. glabra W. A. Rodrigues & H. C. 

Lima, Acta Amazonica 19: 149. 1989 [1990]. Type. Brazil. Amazonas: Mun. Presidente Figueiredo-Balbina, rio 

Uatuma, 25 Sep 1987 (fl., fr.), F. Matos et al. 190 (holotype: 

INPA 148166!; isotypes MG, RB).

Inflorescences glabrous.

FIGURE 18. Geographic distribution of Paloue induta Sandwith subsp. induta (dots) and Paloue induta subsp. glabra W. A. 

Rodrigues & H. C. Lima (triangle) in Guyana and Brazil.
NUMBER 109 • 2 3

Distribution. Endemic to the state of Amazonas, 

Brazil.

Phenology. Collected in flower in September and in 

fruit in March and September.

Notes. The subspecies, as presently circumscribed, is 

restricted to a very specific locality in Amazonas, Brazil (the 

riparian vegetation of the Uatumã and Pitinga Rivers near the 

UHE de Balbina). As for many specimens of Paloue, it is difficult to identify them to species without flowering material, and 

all available specimens examined were in fruit. As described, 

this subspecies differs from P. induta subsp. induta in that 

the inflorescence is glabrous and not covered in light-brown 

trichomes. However, other specimens of P. induta, as well as 

those of P. riparia and P. guianensis, exhibit a gradation in 

trichome density. The presence or absence of trichomes may 

prove to be an unreliable character for species and subspecies 

identification, but until more collections are available, subspecies glabra is retained as a distinct taxon. Unfortunately, herbarium material did not prove suitable for molecular analysis 

that might help verify the relationships among these closely 

related taxa.

Additional specimen examined. BRAZIL. Amazonas: Mun. 

Pres. Figueredo, Rio Uatumã entre rios Pitinga e Uatumã (localidade Nazare), 1–2°S, 59–60°W, 18 Mar 1986 (fr.), Cid Ferreira

et al. 6795 (MO!, NY!).

11. Paloue leiogyne (Ducke) Redden comb. nov.

Elizabetha leiogyne Ducke, Bull. Mus. Nat. Hist. Paris 4: 727. 1932. 

Type. Brazil. Amazonas: São Gabriel, Rio Negro, 30 Nov 1929 (fl.), 

A. Ducke s.n. (lectotype: RB 23285; isolectotypes: G, IAN, K!, NY!, 

R, U!, US!); remaining syntype: Serra Curicuriary, A. Ducke s.n. (RB 

23286).

FIGURES 19, 20

Tree 15–30 m tall, branchlets pilose to tomentose. Stipules caducous, oblanceolate, the apex acute and mucronate, 

5.5–6.5 × 0.5–1.5 cm, glabrous or slightly pubescent near 

the apex or base. Leaves compound, alternate, paripinnate; 

petiole (1–)2.5–3.5 mm long, terete, pilose to tomentose, 

rachis (12–)15–22(–26) cm long, axis pilose, narrowly alate, 

the wing slightly spreading, without an apical projection, 

puberulous, constricted at each leaflet pair insertion; leaflets

37–50(–57)-jugate, sessile, narrowly oblong, 19–28 × 2.5–5 mm, 

the base inequilateral, the apex rounded, sometimes slightly 

retuse, with a small caducous mucro, the margin sometimes 

microscopically ciliolate along one or both sides of the leaflet, a tuft of long hairs usually developed near the base of the 

midvein on the lower surface, midvein salient on both surfaces, the venules more or less prominulous, especially on the 

lower surface. Inflorescences usually terminal racemes, 5–7 cm 

long, the rachis glabrous, or pilosulose near the base or at the 

insertion of the pedicels; bracts rounded, 17–19 × 14–17 mm, 

glabrous or sometimes ciliolate; bracteoles oval, united 

marginally into a tube enclosing the flower bud, glabrous, 

12–18 mm long; pedicels 2–4 mm long, glabrous. Flowers:

“greenish white with very long erect stamens” (fide Ducke, 

1934); the hypanthium glabrous, stipe (3–)5–8.5 mm long, the 

tube 9–12 mm long, 8–10 mm in diameter; sepals pale green 

or the inner surfaces white, glabrous, oblong, (12–)16–17 × 

6–9 mm, rounded to obtuse; petals 5, white, the dorsal one 

14–18 × 10–16 mm, oblong-ovate, bi-auriculate basally, ciliate 

apically and villose at the extreme base on the inner side, the 

other four petals elliptic, ciliate apically, 15–21 × 5–7.5 mm; 

stamens 9, 3 fertile, white, the fertile stamen filaments 20–28 

mm long, villose, joined with the sterile, glabrous filaments 

(these 10–11 mm long) into a tube (except at the insertion of 

the dorsal petals) 3–5 mm long, anthers red, elliptic, 6–7.5 × 

2.5–3 mm; ovary glabrous, 8.5–10 × 2–2.5 mm, narrowly 

oblong, the gynophore glabrous, about 2 mm long, style 14–26 

mm long, glabrous, stigma capitellate. Fruit a legume, oblong, 

ca. 20 × 5 cm, ca. 4–9-seeded.

Distribution. Brazil: Amazonas; Venezuela: Amazonas.

Phenology. Flowering October and November, fruiting October to April.

Note. R. S. Cowan (1976: 336) selected the specimen 

Ducke [s.n., RB] 23285 in RB as the lectotype of Elizabetha leiogyne from among the two syntype collections (RB 23285 and RB 

23286) cited by A. Ducke in the protologue.

Additional specimens examined. VENEZUELA.

Amazonas: Cerro de la Neblina, Rio Yatua, 200 m, 2 Nov 1957 

(fl.), Maguire et al. 41986 (K!, NY!, P!, RB!, US!); Rios Pacimoni-Yutua, Casiquiare, 100–140 m, 7–8 Dec 1957 (fr.), Maguire et al. 36705 (NY!, US!); Rios Pacimoni-Yutua, Casiquiare, 

100–140 m, 25–26 Oct 1957 (fl.), Maguire et al. 41952 (K!, 

NY!, P!, U!, US!); Rios Pacimoni-Yutua, Casiquiare, 100–140 

m, 9 Jan 1958 (fr.), Maguire et al. 42602 (NY!); Departamento 

Rio Negro, from Los Tambores of Comision de Limite to Caño 

Erubichi on Rio Baria, 1°02′N, 66°20′W, 130 m, 2 Apr 1984 

(fr.), Liesner 17125 (NY!); S. Gabriel (Waupé), R. Negro, 19 Feb 

1975 (fr.), Ribeiro 15.286 (713) (NY!); Rio Negro, S. Gabriel, 

Nascimento et al. 148; 198 (NY!).

BRAZIL. Amazonas: Rio Negro, 14 Jan 1933 (sterile), 

Ducke 157 (F!, MAD, NY!); Mun. São Gabriel da Cachoeira, 

Estrada para Camanaus, logo apos o entroncamento da 

Estrada para o aeroporto, 24 Nov 1987 (imm. fr.), de Lima

et al. 3298 (MO!, NY!); Rio Negro: Rio Cauaburi, Maturacá, 

5 Jul–12 Aug 1967 (st.), Schultes 24578 (K! image, U!); Vicinity of Maturacá Mission along Rio Maturacá, 20 Oct 1970 

(fr.); vicinity of Maturaca Mission, also Rio Maturaca, 20 Oct 

1970 (st.), Steyermark 104038 (K!, NY!, US!); Manaus–Caracarí road, ZF-2 road, 50 m off the road, 11 Dec 1980 (st.), 

Renner 59 (US!).

12. Paloue macrostachya (Benth.) Redden comb. nov.

Elizabetha macrostachya Benth., Fl. Bras. 15(2): 214. 1870. Type. Brazil. Gapo of Rio Paapuris, Nov 1852 (fl., fr.), R. Spruce 2671 (holotype: K!).
2 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 19. Paloue leiogyne (Ducke) Redden. Reproduced from Cowan (1976): a, habit; b, portion 

of leaf rachis; c, one bract; d, bracteoles; e, flower at anthesis with bracteoles removed; f, adaxial petal 

(left) and lateral or abaxial petal (right). All drawn from Maguire et al. 41952 by Nancy M. Goor.
NUMBER 109 • 2 5

FIGURE 20. Geographic distribution of Paloue leiogyne (Ducke) 

Redden (dots) in Venezuela and Brazil.

FIGURES 21, 29B

Tree 10–15 m tall, branchlets glabrous. Stipules not seen. 

Leaves compound, alternate, paripinnate; petioles 9–14 mm long, 

terete, usually glabrous but occasionally minutely puberulous, 

rachis 14–28 cm long, terete, without wing or apical projections, 

glabrous or minutely puberulous overall or only at insertion 

of leaflets; leaflets 5–7 jugate, the petiolules 1.5–5 mm long, 

minutely puberulous, blades elliptic, 9.5–14 × 3.5–5 cm, the base 

attenuate, the upper side shorter, the apex abruptly acute to acuminate, glabrous on the upper surface, glabrous and microscopically dotted on lower surface. Inflorescences terminal racemes, 

25–100 cm long, rachis glabrous; bracts 5–8.5 × 3.5–4.5 cm, 

glabrous, rigid-coriaceous, obovate, striate on the inner surface; 

bracteoles 5.0–5.5 × 1.5 cm, rigid-coriaceous; joined marginally 

for about 3⁄4 of their length; pedicels glabrous, 11–22 mm long, 

longer and with a greater diameter in fruit. Flowers: hypanthium 

glabrous, woody, cylindrical, tapered basally, stipe 5–7 mm long, 

the tube 20–27 mm long and about 10 mm in diameter in flower, 

becoming about 40 mm long in fruit; sepals red, 5.5–6.0 × 

1.1–2.0 cm, more or less oblong, glabrous, the apex cucullate, 

the 3 petals 6.5–7.5 × 1.0–1.7 cm, glabrous, oblong, 2 petaloidia 

oblong, 15–17 × 6 mm; stamens 9, exserted, glabrous, the filaments about 4.5–7.5 cm long, joined basally for about 5 mm, the 

anthers linear, about 23 × 2 mm; ovary velvety. Fruit a legume, 

22–28 × 5–6.5 cm, arcuate-oblong, velvety, the dorsal suture 

somewhat marginate, ca. 6–9-seeded.

Distribution. Brazil: Amazonas; Venezuela: Amazonas.

Phenology. Collected in flower in November, February, March, and May, and in fruit in February, April, and May.

Notes. Paloue macrostachya is a dominant tree in 

areas of white sands and black water in Venezuela. These 

areas are interspersed with areas of laterite soil, and the vegetation on the two types of soils is distinctly different. Paloue 

macrostachya only occurs on the white sands. After collecting Redden 3714, the stem was put in a bucket of water to 

preserve the specimen for photography. During the night a 

loud “pop” was heard, which was the sound of the dehiscing woody bracteoles. After bracteole dehiscence, the flower 

slowly emerged.

FIGURE 21. Geographic distribution of Paloue macrostachya

(Benth.) Redden (dots) in Brazil and Venezuela.
2 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Additional specimens examined. BRAZIL. Amazonas: Beira do Rio Icana, catinga alta, 12 May 1948 (fl.), Black

2654 (U!, US!); Rio Negro, alto Icana, vicinity, Serra Tunuhy, 15 

May 1947 (fl.), Fróes 22282 (U!).

VENEZUELA. Amazonas: 3.5 km from Yavita along YavitaMaroa Road (bearing 233°), 2°54′06′′N, 67°27′27′′W, 120 m, 

19 Feb 1998 (fl., fr.), Berry et al. 6603 (MO!: 2 sheets, VEN); 3–5 

km NE of San Carlos de Rio Negro, ca. 20 km S of confluence 

of Rio Negro and Brazo Casiquiare; 1°56′N, 67°03′W, 120 m, 

1 May 1979 (fl.), Liesner 7128 (MO!); 3–5 km NE of San Carlos de Rio Negro, ca. 20 km S of confluence of Rio Negro and 

Brazo Casiquiare; 1°56′N, 67°03′W, 120 m, 4 May 1979 (fl., fr.), 

Liesner 7200 (MO!); Entre San Carlos de Rio Negro y El Solano, 

Departamento Río Negro, 11–17 Mar 1979 (fl.), MarcanoBerti & Salcedo 67-979 (US!); road between San Carlos and 

Solano, 1°55′48.8′′N, 67°00′58.8′′W, 101 m, 15 Feb 2005 (fl., 

imm. fr.), Redden 3714 (K!, NY!, PORT, US!, VEN); 7–9 km 

de Yavita hacia Pimichin, 2°55′N, 67°25–30′W, 125 m, 22 Apr 

1970 (fr.), Steyermark & Bunting 102923 (US!, VEN!).

13. Paloue paraensis (Ducke) Redden comb. nov.

Elizabetha paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 3:102. 1922. 

Type. Brazil. Pará: Cachoeira do Mangabal, Rio Tapajoz, 4 Sep 1916, 

A. Ducke 16449 (lectotype: MG, isolectotypes: G, P!, US!).

FIGURE 22

Large tree 16–30 m tall, 20 cm or more in diameter, the 

brachlets pilose and puberulous, the hairs more or less retrorse. 

Stipules 42–60 × 10–22 mm, oblanceolate, slightly bifid, thin, 

puberulous externally, glabrous within, caducous. Leaves compound, alternate, paripinnate; petioles 2.5–3 mm long, pilose and 

puberulous, rachis 9–16.5 cm long, pilosulose, narrowly winged, 

wing puberulous or glabrous with apical projections at leaflet 

insertions; leaflets sessile, 32–38-jugate, 17–27 × 3–5.5 mm, 

oblong, base inequilateral, apex truncate to rounded, retuse and 

mucronulate, the mucro caducous, glabrous except for a pilose 

patch at the base of one side of the salient midvein, venules subobscure. Inflorescences terminal, sessile, compact racemes, 4–8 cm 

long, the axis strigose-sericeous; bracts caducous, oval, cucullate, 15–27 × 8–15 mm, puberulous externally, glabrous within, 

ciliate; bracteoles 14–19 mm long, united marginally into a tube 

enclosing the flower bud, puberulous externally; pedicels 2–4 mm 

long, puberulous, hairs often retrorse. Flowers: hypanthium 

puberulous or pilosulose, stipe (6–)10–15 mm long, the cupulate 

tube 3–7 mm wide and deep; sepals white, the dorsal one 11–17 × 

8–12 mm, oblong-oval to rotund, the other 4 oblong, 12–18 × 

5–9 mm, minutely puberulous externally; petals 5, white, retuse 

and ciliate apically, the dorsal one triangular-ovate, 10–18 × 6–8 

mm, the other four 12–18 × 4–7 mm; stamens 9, white, 3 fertile, 

the filaments connate in a short tube except at the adaxial petal, 

those of the fertile stamens 50–58 × 1.5–2 mm, glabrous, the 

staminodia ca. 15 mm long, glabrous; ovary narrowly oblong, 

velutinous, 7–10 × 1.3 mm, 8-ovulate, the stipe 5–10 mm long, 

sub-sigmoid, glabrous except puberulous apically, style 34–40 

mm long, glabrous, stigma capitate. Fruit a legume, 8–11 × 

2.5–4 cm, ca. 4–6-seeded.

Distribution. French Guiana; Suriname; Brazil: 

Amapa, Pará.

Phenology. Collected in flower in September, October, 

and January and in fruit in January.

Note. Ducke 16449, a sterile collection, was the one 

chosen as the type of P. paraensis by Ducke (1934) in his later 

reconsideration of his own species (Cowan, 1976).

Additional specimens examined. FRENCH GUIANA. Cayenne: Trois Sauts, sous le village Pina, 11 Jan 1975 (fl., 

imm. fr.), Grenand 649 (US!).

SURINAME. Brokopondo: Nassau Mountains, Marowijne 

River, 525 m, 4 Mar 1955 (st.), Maguire 40752 (NY!).

BRAZIL. Amapa: Rio Araguari, vicinity camp 12, 1°11′N, 

52°08′W, 29 Sep 1961 (st.), Pires et al. 51310 (NY!, US!); Pará: 

Belém in horto Musei culta, e regione medii fluminis Tapajoz 

(Igarape das Pedras, prope cataractam Furnas) introducta, 

FIGURE 22. Geographic distribution of Paloue paraensis

(Ducke) Redden (dots) in Brazil.
NUMBER 109 • 2 7

21 Oct 1940 (fl.), Ducke 591 (MO!, NY!, US!); Rio Itapacura 

afl. do Tapajoz, 24 Jan 1933 (st.), Ducke 23723 (K!, NY!); upper 

Cupary River, plateau between the Xingu and Tapajos Rivers, 

Sep 1931 (fl.), Krukoff 1167 (K!, NY!); upper Cupary River, 

plateau between the Xingu and Tapajos Rivers, Sep 1931 (st.), 

Krukoff 1184 (K!, NY!); Belem do Pará, Museu Goeldi, culta 

proced. Igarapi dos pedras prope Furnas, R. Tapajoz, 1936 (jv. 

fr.), Sigueira 34958 (K!, US!); Jardim Botânico Rio de Janeiro, 

7 May 1984 (st.), de Lima 2156A (K!, NY!).

14. Paloue princeps (M. R. Schomb. ex Benth.) Redden comb. 

nov.

Elizabetha princeps M. R. Schomb. ex Benth., Hook. Jour. Bot. 2: 92. 

1840. Type. Guyana. 1839 (fl., fr.). R. Schomburgk s.n. (holotype: 

K!; isotype: F!).

FIGURE 23

Medium-sized to large tree (8–)20–35(–50) m tall, trunk 

(25–)40–50 cm in diameter, the outer bark greenish gray, smooth, 

thin, the inner bark brown, branchlets pilose or pilosulose, glabrescent. Stipules caducous, (3.5–)4.5–9.5 × (1.1–)1.5–3.0 cm, 

narrowly to broadly oblanceolate, the apex rounded, entire, thin, 

densely puberulous on the outer surface, glabrous on the inner 

surface, often ciliolate on margins. Leaves compound, alternate, 

paripinnate, oblong–lanceolate; the petioles 1.5–4 mm long, 

terete, more or less pilosulose, occasionally glabrous, rachis (12–)

20–25(–30.5) cm long, pilosulose, the narrow wing erect with 

apical projections at leaflet insertions, usually glabrous; leaflets

(27–)32–40(–51)-jugate, sessile, narrowly oblong, (18–)30–47 × 

(3–)5–7(–8) mm, base inequilateral to obtuse, apex tapering but 

rounded to obtuse, glabrous except for a patch of long hairs near 

the base on one side of the midvein on the leaflet undersurface, 

often completely glabrous at maturity, micro-punctate on the 

lower surface, the venation sub-obscure or the midvein more 

or less salient. Inflorescences dense, strobiliform racemes, sessile, axillary or terminal, 6–11 (–15) cm long, the rachis densely 

pilose or strigose; bracts usually sub-rotund, infrequently oval or 

broadly ovate, (25–)28–3 × (22–)24–35 mm, cucullate, rounded 

to obtuse apically, coriaceous, pilose or pilosulose externally, glabrous within, ciliate or ciliolate; bracteoles united for about 2⁄3 of 

their length, acute, (20–)23–30 mm long, united marginally into 

a tube enclosing the flower bud, externally pilose and pilosulose, the inner surfaces pilose, tomentose, or strigose, at least in 

part; pedicels (2–)5–8 mm long, pilosulose, pilose, or tomentose. 

Flowers: hypanthium pilosulose or pilose, stipe (9–)15–20 mm 

long, the tube cupular, 5–11 mm long, 7–11 mm in diameter; 

sepals 4, white to cream-colored, 13–24 × 4–15 mm (the dorsal 

one broader), elliptic–oblong, oval or oblong, pilosulose externally, glabrous within; petals 5, white, except for the extreme 

base, which is red, the dorsal one triangular-ovate, the apex ciliate, usually emarginate, base obtuse to cordate, 1.2–2.0(–2.4) × 

0.9–1.5(–2.4) cm, the other 4 petals elliptic to oval or oblong, 

1.8–2.7 × 0.5–1.2 cm, glabrous; stamens 9, red, 3 fertile ones 

with larger filaments, 3.0–5.8 cm long, villose at least basally, 

the sterile filaments glabrous, slender, 1.0–2.5 cm long, united 

basally with the fertile ones in a glabrous tube 0.5–1.5 mm 

long, anthers narrowly elliptic to oblong, 7–11 × 1.5–3 mm, 

glabrous, pollen grains 57–60 μm in diameter, uniformly verruculose; the ovary narrowly oblong, velutinous, 7.5–16 × 2–3.5 

mm, 10–12-ovulate, the stipe (2.5–)5–8(–10) mm long, glabrous, 

style 2.3–4.0 cm long, glabrous, stigma capitate. Fruit an oblong 

legume, sometimes slightly arcuate, flat, narrowly bi-alate dorsally, velutinous, 17–35 × 4–6.2 cm; ca. 4–6-seeded.

Distribution. Guyana; French Guiana; Suriname; Brazil: Amazonas, Roraima; Venezuela: Amazonas.

Phenology. Flowering from September to May, fruiting from September to April and in August.

Additional specimens examined. VENEZUELA. 

Amazonas: Atabapo, alto Rio Orinoco, bosques riberenos 

medios en la planicie alluvial, 30 km al SE de le Esmeralda, 19 

Feb 1990 (fl.), Aymard 8029 (NY!); Atabapo, alto Rio Orinoco, 

bosques medios a altos sobre planicie alluvial, 35 km al SE de 

FIGURE 23. Geographic distribution of Paloue princeps (M. R. 

Schomb. ex Benth.) Redden (dots) across the Guiana Shield.
2 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

la Esmeralda, ca. 2°58′N, 65°21′W, 180 m, 16 Feb 1990 (st.), 

Aymard & Delgado 7887 (NY!); Rio Orinoco below Raudal y 

Salto El Tobogan, 2°14′N, 63°45′W, 370 m, 21 Nov 1989 (fl.), 

Berry 4797 (K!, MO!); Rio Mawarinuma, Neblina Base Camp, 

0°50′N, 66°10′W, 140 m, 9 Feb 1985 (fr.), Boom & Weitzman 

5718 (NY!, U!); Ugueto, 30 Sep 1951 (fl., fr.), Croizat 766

(NY!); without precise locality, 1951 (fl.), Croizat 925 (NY!); 

Atabapo, Salto Yureba, Cerro Yureba, lower Ventuari, 4°03′N, 

66°01′W, 350 m, 15–16 Mar 1985 (fl., jv. fr.), Liesner 18701

(MO!); Orillas rio Tucano, aflueute del Cauaburi, 100 m, 16 

Apr 1964 (fr.), Ewel 104 (NY!); Cerro de la Neblina, Rio Yatua, 

100–140 m, 7–8 Dec 1953 (st.), Maguire et al. 36736 (F!, NY!, 

US!); Cerro de la Neblina, Río Yatua, 140 m, 14 Dec 1953 (fl.), 

Maguire et al. 36796 (F!, K!, NY!, US!); 1.5 km S of base camp, 

SE side of Cerro de la Neblina, 00°49′20′′N, 66°09′45′′W, 150 

m, 12 Feb 1985 (fr.), Nee 30831 (NY!); Rio Negro, along Rio 

Baria (=Rio Mawarinuma), just upstream from base camp on 

SW side of Cerro de la Neblina, 00°49′50′′N, 66°09′40′′W, 140 

m, 27 Jan 1985 (fr.), Nee 30573 (NY!); Rio Yatuá, 1°10′30.3′′N, 

66°3′25.8′′W, 105 m, 4 Feb 2005 (fr.), Redden 3536 (PORT!, 

US!, VEN!); Missão Salesiana de Maturaca, R. Maturaca, afl. do 

R. Cauaburi, without date (st.), Souza s.n. (NY!); Rio Metacuni, 

Selvas ribereñas abajo del Salto Wânitima, en el Caño Guacaguaca, 180–200 m, 21 Jan 1990 (fr.), Stergios & Velazco 14003

(MO!, NY!, PORT!); Rio Metacuni, Tepuy-Altiplanicie Sedukerawa y areas adyacentes, 3°15′N, 64°56′W, 210–650 m, 26 

Jan 1990 (fr.), Stergios & Velazco 14256 (MO!, PORT!); Mission Rio Mavaca, 1°58′N, 65°06′W, 200–230 m, 5 Feb 1991 

(fr.), Sterigos & Yanez 15203 (NY!, PORT!); Rio Marawinuma, 

vicinity of Base Camp, 0°50′N, 66°10′W, 140 m, 19 Apr 1984, 

Thomas & Plowman 3151 (NY!); along Rio Marawinuma, SE 

of base camp, 0°50′N, 66°09′W, 140 m, 30 Apr 1984, Thomas 

& Samuels 3290 (NY!); Lower Rio Guainia, 100 m, 1942 (st.), 

Williams 14790 (F!, G, NY!, US!).

GUYANA. Potaro-Siparuni: Iwokrama Rainforest Reserve, 

Iworkrama Mts., 0–1 km SE of camp at bottom of gorge, 4°20′N, 

58°48′W, 75–150 m, 24 Mar 1997 (fr.), Clarke 4202 (BRG!, 

NY!, US!); Iwokrama Mts., base camp, 4.6 km E of Georgetown–Lethem Road, Plot 1-1, 04°20′N, 58°48′W, 70–150 m, 

22 Nov 1995 (st.), Hoffman 4877 (BRG!, NY!, US); Iwokrama 

Rainforest Reserve, Pakatau Falls area, old trail to Mahdia, 

4°45′9′′N, 59°1′21.2′′W, 65 m, 30 Oct 2002 (st.), Redden 1104

(BRG!, K!, MO!, NY!, US!); Upper Takutu, cut line GH from 

Lethem to Kurupukari Road to Iwokrama Mts., 4°19′02.77′′N, 

58°52′01.61′′W, ca. 200 m, 30 Apr 1992 (st.), Pennington & 

Johnson 431 (K!); U. Demerara-Berbice Region. Purburi Forest 

Reserve Camp, 5°1′48.5′′N, 58°36′41.9′′W, 55 m, 23 Oct 2002 

(fr.), Redden et al. 1080 (K!).

SURINAME. Brokopono: In montibus, qui docuntur Nassau, 19 Mar 1949, Lanjouw & Lindeman 2851 (NY!). Sipaliwini: Tumuc Humac Mts., Talouakem, second base camp, 

2°29′N, 54°45′W, 300 m, 15 Aug 1993 (fr.), Acevedo-Rodriguez 

6044 (NY!); along Ulemari River, ca. 150 km upstream from 

its confluence with Litani River, 2°46′5′′N, 54°51′15′′W, 175 m, 

19 Apr 1998 (fr.), Hammel et al. 21584 (K!, MO!, US!); Upper 

Suriname River, Gran Lio, Awarradan vicinity, 1.5 km at 170° 

from tourist camp island (access from Sandu Matu Creek trailhead, 

3°50′00.57′′N, 55°36′51.98′′W) (B. Hoffman 1.0 ha research plot 

ST2), 3°49′56.14′′N, 55°36′39.53′′W, 22 Nov 2005 (st.), Hoffman 6635 (US!); same locality, 19 Mar 2006 (st.), Hoffman 6794

(US!); Wilhelmina Gebergte, 1.5 km southeast of Juliana Top, near 

headwaters of West Rivier, 3°36′–3°41′N, 56°30′–56°34′W, 750 

m, 26 Aug 1963 (fr.), Irwin et al. 55072 (NY!, U!).

FRENCH GUIANA. Cayenne: Saut Parasol-Bassin du Sinnamary, 4°20′N, 52°54′W, 65 m, 11 Jan 1992 (fl.), Cremers 

7323 (AAU, B, BM!, BR, CAY!, G, K!, MO!, NY!, P, U!, US!). 

Saint Laurent du Maroni: Saül, Monts La Fumée, 3°37′N, 

53°12′W, 200–400 m, 9 Oct 1982 (st.), Boom & Mori 1932

(NY!); Saül, Monts La Fumée, 3°37′N, 53°12′W, 200–400 m, 

9 Oct 1982 (st.), Boom & Mori 1940 (NY!); Saül, Monts 

La Fumée, 3°37′N, 53°12′W, 200–400 m, 9 Oct 1982 (st.), 

Boom & Mori 1950 (NY!); Saül, Monts La Fumée, 3°37′N, 

53°12′W, 200–400 m, 9 Oct 1982 (st.), Boom & Mori 1953

(NY!); Saül, Monts La Fumée, 3°37′N, 53°12′W, 200–400 

m, 9 Oct 1982 (st.), Boom & Mori 1958 (NY!); Saül, Monts 

La Fumée, 3°37′N, 53°12′W, 200–400 m, 9 Oct 1982 (st.), 

Boom & Mori 1960 (NY!); Saut Parasol, Bassin du Sinnamary, 

4°20′N, 52°54′W, 65 m, 11 Jan 1992 (fl.), Hoff 7323 (K!); Saül 

and vicinity, forest camp ca. 10 km ESE from Eaux Claires, 

3°37′N, 53°12′W, 314 m, 4 Sep 1994 (fl.), Mori et al. 23777

(NY!); Nouragues Field Station and vicinity between Camp 

Inselberg and Saut Pararé (all collections made from 1,500–

3,000 m S of Camp Inselberg), 4°05.289′N, 52°40.774′W, 

23 Feb 2002 (fr.), Mori et al. 25433 (K!, NY!).

BRAZIL. Roraima: SEMA Ecological Reserve, Ilha de 

Maracá, Rio Puruí, 3°13′N, 62°31′W, 25 May 1987 (fl.), Milliken & Bowles 290 (K!, NY!); Posto Mucajai, Rio Mucajai, 

vicinity of Mucajai airstrip, 13 Mar 1971 (fr.), Prance et al. 

10936 (NY!, U!, US!); Indian trail from Surucucu near Maitá 

Indian village, 3°20′N, 63°24′W, 14 Feb 1971 (fr.), Prance

et al. 10517 (K!, NY!, U!); Indian trail from Surucucu, 

between Maitá Indian village, 3°20′N, 63°24′W, and Paramiteri Indian village, 3°25′N, 63°03′W, 20 m, 20 Feb 1971 (fr.), 

Prance et al. 10639 (K!, NY!, U!); Rio Cauaburí, between 

Anta and Palmito camps, 600–1,300 m, 16 Dec 1965 (fl.), 

Silva & Brazão 60669 (K!, NY!). Amazonas: Serras do Jacamin, Rio Negro, 5 Dec 1936 (fl.), Ducke 302 (F!, MAD, NY!), 

5 Dec 1936 (fl., fr.), Ducke 29025 (K, 2 sheets!); Silva in declivibus montium Jacamin, Rio Negro super Santa Izabel, 5 Dec 

1936 (fl.), Ducke 328 (NY!); Rio Curicuriary, 17 Nov 1936 

(fl.), Ducke 34962 (K!).

15. Paloue riparia Pulle, Enum. Vasc. Pl. Surinam 212, pl. 10. 

1906. Type. Guyana (as British Guiana). Saramacca River, 

near Janbasigado, Jan 1903 (fl.), A. Pulle 154 (holotype: U!; 

isotype: U!).
NUMBER 109 • 2 9

FIGURES 24, 25, 30B

Small to medium-sized tree, 7–10 m tall, branchlets gray–

brown, glabrescent. Stipules caducous, not seen. Leaves simple, 

alternate, entire, ovate to oblong, 9–16 × 2.5–7 cm, apex acuminate, base rounded to acute with crater-like glands at the base 

of the lamina, glabrous; petioles 3–7 mm long. Inflorescences

umbel-like, terminal racemes, 7–12 cm long, minutely puberulous 

or glabrate; bracts orbiculate, 5 mm long, caducous; bracteoles 

connate, 5 mm long, bilobed, rounded, persistent; pedicels 7–12 

mm long, angular. Flowers: hypanthium 15 mm long including 

the stipe; sepals deep red, 9–11 × 6 mm, rounded at the apex, 

reflexed; petals 5, red, largest petal 5–7 × 3 mm, the other 4 petals very small, 1–3 × ca. 1 mm, margins pilose; stamens 9, red, 

filaments 5–6 cm long, connate unequally to form a tube, longpilose; anthers oblong, glabrous, 5 mm long, , ovary 1 cm long, 

glabrous, style 2 cm long, stigma capitate. Fruit a sub-falcate 

legume, 11–16 × 2.5–4 cm, stipe to 2.4 cm long and 1–2 mm 

thick, glabrous, apex obliquely acuminate, ca. 6–8-seeded.

Distribution. Guyana, French Guiana, Suriname.

Phenology. Flowering from September to April, fruiting from September to June.

Notes. This species can be distinguished from the others 

in the genus by the small bracteoles and flowers and the glabrous 

anthers and gynoecium. The flowers have only one large petal 

and four small ones.

On numerous occasions, hummingbirds were observed feeding in the flowers of this species. The cup-shaped hypanthium 

was filled with liquid. Small holes at the base of the hypanthium 

also were observed, which may have been produced by “nectarrobbing” birds. Comparable small holes have not been observed 

on herbarium specimens.

Additional specimens examined. GUYANA.

Cuyuni-Mazaruni: Upper Mazarooni, Appun, 1702 (K!); 

Appun, 263 (K!); Takutu Creek to Puruni River, Mazaruni River, 

8 Dec 1944 (fl., im. fr.), Fanshawe 4912 (K, 2 sheets!, NY!, U!); 

Fanshawe 4881 (F!, K, 2 sheets!, NY!, U!); Eping River, Lang 

413 (NY!); Lang et al. 292 (F!); McDowell 3758 (BRG!, K!, 

MO!, NY!, US!); Kamakusa, Lang et al. 332 (F!, NY!, US!); 

Kuyuwini River, first base camp, 2°4′0.0′′N, 59°09′32.8′′W, 273 

m, 21 Apr 2004 (fl., fr.), Redden 3106 (BRG!, K!, NY!, US!); 

Kamakusa, Mazaruni River, 5°56′10.3′′N, 59°54′6.5′′W, 64 m, 

29 Sep 2004 (fl., fr.), Redden 3208 (BRG!, K!, NY!, US!), Mazaruni River, small rock and sand islands in river, 6°19′30.7′′N, 

58°42′20.3′′W, 2 m, 18 Oct 2004 (fl.), Redden 3314 (BRG!, 

K!, NY!, U!, US!); Mazaruni River, 0.22 mi E of base camp, 

5°53′1.2′′N, 59°52′51.9′′W, 89 m, 4 Oct 2004 (fl.), Redden 

3229 (BRG!, K!, NY!, U!, US!); Mazaruni River, 0.67 mi SW of 

base camp, 5°48′46.4′′N, 59°35′7.8′′W, 64 m, 6 Oct 2004 (fl., 

fr.), Redden 3233 (BRG!, K!, NY!, U!, US!); Mazaruni River, 

unnamed falls (possibly Carin Crow Falls) 0.31 mi W of base 

camp, 5°57′17.1′′N, 59°18′48.1′′W, 58 m, 9 Oct 2004 (fl., imm. 

fr.), Redden 3263 (BRG!, K!, NY!, U!, US!); Mazaruni River, 

small islands along river, 1.34 mi E of base camp, 5°59′52.6′′N, 

59°12′44.4′′W, 63 m, 11 Oct 2004 (fl.), Redden 3281 (BRG!, 

K!, NY!, U!, US!); Mazaruni River, 0.78 mi NE of base camp, 

6°6′34.4′′N, 58°54′6.6′′W, 21 m, 14 Oct 2004 (fl., imm. fr.), Redden 3305 (BRG!, K!, NY!, U!, US!). Potaro-Siparuni: Siparuni 

River, Pakutau Falls; 350 m along survey line due S from falls, 

4°45′N, 59°1′W, 26 Apr 1992 (fr.), Hoffman et al. 1454 (BRG!, 

photo at K!, NY!, U!, US!); Iwokrama Rainforest Reserve, 5 km 

S of Siparuni River, Pakutau Plot 1, 4°45′17.2′′N, 59°1′27.8′′W, 

8 Nov 1995 (st.), Hoffman et al. 4759 (BRG!, US!); Iwokrama, 

Siparuni River, Pakutau Falls, 5 km upstream Mutchnick 520

(BRG!, NY!, US!); main trail, swampy area, 0.3 mi E of Whitewater base camp, 4°43′54.2′′N, 58°50′43.1′′W, 115 m, 6 Nov 

2002 (fl.), Redden 1161a (BRG!, K!, NY!, US!). Upper Demerara-Berbice: Mabura region, Central Demerara compartment, 

main road to Pibiri, 5°20′N, 58°30′W, 30 Nov 1992 (fl.), Ek

et al. 614 (NY!, U!).

SURINAME. Nickerie: Vicinity of Blanche Marie Waterfall on the Nickerie River, 4°45′30′′N, 56°52′50′′W, 50 m, 

25 Jul 1995 (fr.), Evans et al. 2327 (MO!, RB-digital photo, 

US!); 19 Nov 1995 (fl.), Evans et al. 2446 (K!, MO!, US!). 

Sipaliwini: N side of Kurini Island, E of trail from airstrip to 

N branch of Kuruni River around island, ca. 30 airline km 

E of confluence on Corantijn River, 3°22′12′′N, 57°20′36′′W, 

180 m, 8 Nov 1994 (fl.), Evans et al. 1922 (CAY!, K!, MO!, 

RB-digital photo, U!, US!); Coppename River, Raleigh Falls, 

Sept 1933 (fl., imm. fr.), Lanjouw 732 (L!, U!); Oever Coppename River boven Raleighvallen, 27 Nov 1954 (fl.), Mennega 

548 (NY!, U!); Central Suriname Nature Reserve, Coppename River, Tonckens Falls, 4°25′21.7′′N, 56°31′34.2′′W, 

60–90 m, 23 Feb 2004 (fr.) Clarke 11061 (US!); Central 

Suriname Nature Reserve, lower Rechter Coppename River, 

4°18′9′′N, 56°27′13.7′′W, 60–90 m, 24 Feb 2004 (fr.), Clarke 

11094 (US!); Central Suriname Nature Reserve, back channel of Coppename River at ‘Sidonkroetoe Sula’ (= sit down 

meeting rapids), 4°32′14.9′′N, 56°30′52.5′′W, 60–90 m, 9 

Mar 2004 (imm. fr.), Clarke 11338 (US!); Central Suriname 

Nature Reserve, unnamed tributary of Coppename River, 

200 m south of Driefoetoe Sula (= 3-ft rapids), 4°36′41.9′′N, 

56°30′24.6′′W, 60–90 m, 10 Mar 2004 (imm. fr.), Clarke 

11374 (US!); Upper Tapanahoni River, mouth of Paloemeu 

River, 11 Feb 1936 (fl.), Rombouts 674 (K!, U!, US!); Natuurreservaat Raleighvallen-Voltzberg op rivieroever nabij 

GMD-kamp, 1 Nov 1972 (fl.), Teunissen & Schulz 13650 (K! 

image of sheet at U, NY!, U!). Sipaliwini: Vicinity of camp on 

W bank of Zuid River, across river from (i.e., W and outside 

of) Central Suriname Nature Reserve, ca. 10 km straight-line 

distance SSE of Kayserberg Airstrip, 3°00′03′′N, 56°27′03′′W, 

240 m, 14 Jun 2003 (fr.), Evans et al. 3516 (MO!, US!); 

large island in Litani River, 6 km upstream from its confluence with Marowini River to form Lawa River, 3°17′22′′N, 

54°06′52′′W, 150 m, 1 Apr 1998 (fr.), Hammel et al. 21242

(K!, MO!, US!).
3 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 24. Paloue riparia Pulle. Reproduced from Pulle (1906: pl. 10, p. 212).
NUMBER 109 • 3 1

FIGURE 25. Geographic distribution of Paloue riparia Pulle 

(dots) across the Guiana Shield.

FRENCH GUIANA. Cayenne: Montagne de Kaw sommet, 

300 m, 24 Jan 2000 (fl.) Bordenave & Raes 5135 (CAY!, P, U!); 

Montagne de Kaw, 220–250 m, 12 Dec 1954 (fl.), Cowan 38737

(NY!, RB-digital photo, U!); Cowan 38814 (F!, K!, NY!, US!); 

Campoi: Trois Sauts, 13 May 1976 (fr.), Grenand 1285 (CAY!); 

Pasisit, bord de l’Oyapock, Trois Sauts, 6 Jan 1975 (fl., fr.), Grenand 633 (CAY!, US!).

16. Paloue sandwithii Redden. Brittonia (60) 3: 257–260. 

2008. Type. Guyana. Potaro River below Waratuk Falls, 

05°16′26.4′′N, 59°22′46.7′′W, 72 m, 5 Sep 2006 (fl.), K. M. 

Redden 4022 (holotype: BRG!; isotypes: CAY!, K!, MO!, 

NY!, US!).

FIGURES 26, 27, 29F

Small tree, 4–5 m tall, 20–25 cm in diameter, branchlets 

dark brown with waxy cuticle, glabrous. Stipules intrapetiolar, 

membranous, connate basally for at least 3 mm, distal portion 

of stipule linear, 1.1–1.4 × 0.4–0.6 cm, caducous. Leaves simple, 

alternate, entire, coriaceous, dark green when fresh, elliptic–

oblong, 18.5–32 × 7–11.5 cm, petiole terete, 0.5–0.7 cm in 

diameter, base of leaf acute-cuneate, a pair of crater-like glands 

at either side of the midvein at the base of the lamina, apex acuminate, 1.2–2 cm long, mucronate, venation brochidodromous, 

prominent especially on the lower surface. Inflorescences axillary 

and/or terminal, compact racemes, 0.5–1.5 cm long; bracts that 

subtend the inflorescence persistent, 5–6 × 4–5 mm, green, triangular, acute, strigulose abaxially and the margins ciliolate; floral 

bract 11 × 10 mm, free, ciliolate on margins but otherwise glabrous; bracteoles united almost to apex, enclosing bud except at 

apex, 7–9 × 6–8 mm, green, quadrangular, margins hyaline, membranous, glabrous; pedicel 4 × 2 mm, quadrangular, ca. 9 mm 

from insertion of bracteoles to base of hypanthium. Flowers:

hypanthium cupular, 8 × 6 mm, glabrous; sepals 4, the adaxial 

sepal 12 mm long, the others 15–17 × 6–10 mm, green externally 

with red margins, red internally; petals 5, lateral petals 16–17 × 

4–6 mm, when fresh red with white streaks, adaxial petal 18–19 × 

7 mm, clawed, pale red, glabrous except for the ciliate apex, 

edges crispate stamens 9, all fertile, 4–6 cm long, bright red, 

filaments pilose from base to apex, all united to form a tube 

approximately 1 mm long, anthers 7–8 × 2–3 mm, red, versatile, 

densely pilose along outer edge, elliptic, pollen exine verrucate; 

gynoecium glabrous, 9 × 2 mm at anthesis, stipe 2 mm long, glabrous, style 2.8–4 cm long, stigma capitate, minutely papillose. 

Mature fruit not seen.

Distribution. Endemic to Guyana.

Phenology. Collected in flower in August and September; mature fruits unknown.

Notes. Paloue sandwithii most resembles P. guianensis except it differs in having five subequal petals and large 

coriaceous leaves with prominent venation on their lower 

surfaces.

Additional specimens examined. GUYANA.

Below Kaieteur, Potaro River, Sep–Oct 1881, Jenman 893

(BRG!, K!); Kaieteur trail, 1 Aug 1958 (fl.), V. Graham 197 (K!).

17. Paloue speciosa (Ducke) Redden comb. nov.

Elizabetha speciosa Ducke, Trop. Woods 37: 21. 1934. Type. Brazil. Amazonas: Manaus, 2 Oct 1932 (fl., fr.), A. Ducke 23730 (holotype: RB; 

isotypes: F!, G, K!, MAD, NY!, P!, R, U!, US!).

FIGURE 28

Shrub or small tree to 6 m tall, branchlets pilosulose, glabrescent. Stipules caducous, thin, foliaceous, oblanceolate to 

obovate, 4.5 × 0.9–2.2 cm, the apex bifid, glabrous except near 

the apex puberulous. Leaves compound, alternate, paripinnate; 

petioles 2.5–4 mm long, erect, pilosulose, the rachis 10.5–17 cm 

long, narrowly winged with apical projections at each leaflet 

pair insertion, glabrous above, pilosulose below, rachis pilosulose beneath; leaflets sessile, glabrous except for the obscurely 

ciliolate margin, 22–34-jugate, narrowly oblong, somewhat 
3 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 26. Paloue sandwithii Redden: A, flowering branch; B, adaxial surface of leaf showing 

detail of brochidodromous venation; C, paired crater-like glands on the base of the leaf lamina 

undersurface; D, pair of fused, intrapetiolar stipules; E, stamen filament trichomes; F, distal portion of stamen showing trichomes concentrated along the margins of the anther; G, ovary attachment on adaxial wall of hypanthium. All drawn from the holotype, Redden 4022 (BRG), by Alice 

Tangerini, Smithsonian Institution.
NUMBER 109 • 3 3

FIGURE 27. Geographic distribution of Paloue sandwithii Redden (dots) in Guyana.FIGURE 28. Geographic distribution of Paloue speciosa (Ducke) 

Redden (dots) in Brazil.

arcuate, 22–33 × 5–6 mm, base inequilateral, apex truncate, 

slightly retuse, minutely mucronulate, the mucro caducous, ciliolate along upper margin near base of leaflets, the venation subobscure. Inflorescences sessile, terminal racemes, (2.2–)3.0–6.0 

cm long, glabrous; bracts 10 × 6–9 mm, broadly ovate, glabrous, 

strongly cucullate; bracteoles 10–12 × 5–8 mm, united marginally into a tube enclosing the flower bud, glabrous, somewhat 

carinate, oval, obtuse; pedicels 2.5–5 mm long, glabrous, more 

or less compressed with the bracteole extending the length of the 

pedicel. Flowers: hypanthium glabrous, 9–16 mm long, including the stipe, 5–8 mm in diameter; sepals 4, 10–14 × 5–7 mm, 

glabrous, oblong, apex rounded; petals 5, red, more or less elliptic, 10–12 × 4–5 mm; stamens 9, red, 3 fertile, filaments of the 

3 fertile ones 1.9–2.2 cm long, sparingly villose, the sterile ones 

reduced to minute staminodia, all filaments united basally into a 

short tube to 1 mm long; anthers narrowly elliptic, 6.5–9 × 1.5–2 

mm; ovary narrowly oblong, velutinous, 6–12 × 2–3 mm, the 

stipe 1–1.5 mm long, glabrous except at extreme apex, style glabrous, 12–14 mm long, stigma capitellate. Fruit a legume, 11–13 × 

3–4 cm, 3–7-seeded.

Distribution. Endemic to the state of Amazonas, 

Brazil.

Phenology. Collected in flower in October and in fruit 

in October and November.

Additional specimens examined. BRAZIL. Amazonas: Manaus (Manoas), loco Estrada do Aleixo, silva non 

inundabili, 17 Oct 1935 (fl.), Ducke 48 (F!, K!, NY!, R, US!); 

Reserva Florestal Ducke, Manaus-Itacoatiara, km 26, 2°53′S, 

59°58′W, 8 Nov 1995 (fr.), Sothers et al. 672 (K!, NY!).

Acknowledgments

We thank numerous collaborators for their ideas, discussions, expertise, and observations, in particular, Anne 

Bruneau, Hannah Banks, Vicki Funk, Carol Kelloff, Aleksander Radosavljevic, and Heather Lindon. We thank Lucy 

Smith for the elegant figures of Paloue duckei, P. durissima, 

and P. fanshawei; Sara Alexander for help with the distribution maps; and the reviewers and editors for their constructive 

comments on an earlier version of this manuscript. We also 
3 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 29. A. Paloue durissima (Ducke) Redden (photograph by K. Redden). B. Paloue macrostachya (Benth.) 

Redden (photograph by K. Redden). C. Paloue brasiliensis Ducke (photograph by B. Torke). D. Paloue emarginata

(R. S. Cowan) Redden (photograph by B. Hoffman, for The Amazon Conservation Team). E. Paloue guianensis Aubl. 

(photograph by P. Acevedo, for Smithsonian Institution, Department of Botany). F. Paloue sandwithii Redden (photograph by K. Wurdack, for Smithsonian Institution, Department of Botany).
NUMBER 109 • 3 5

FIGURE 30. A. Paloue induta Sandwith subsp. induta (photograph by K. Redden). B. Paloue riparia Pulle (photograph by K. Redden). C. Paloue coccinea M. R. Schomb. ex Benth.) Redden (photograph by K. Redden). D. Paloue 

grahamiae (R. S. Cowan) Redden (photograph by T. McDowell).

thank the herbaria that provided access to collections and 

facilitated loans, especially BM, F, K, MO, NY, P, U, US, 

and WAG. We extend thanks to the governments, agencies, and 

universities of Guyana, French Guiana, and Venezuela for 

facilitating author K. Redden’s fieldwork in those countries. 

Redden also thanks her Amerindian guides and counterparts 

who spent months in the field with her, especially Claudius 

Perry, Dave Singh, Romeo Williams, and Paul Benjamin. 

This work was supported by the National Science Foundation (Dissertation Improvement Grants DEB-0408041 and 

DEB-0316375), Smithsonian Institution’s Biodiversity of the 

Guianas Program, the New York Botanical Garden (Rupert 

Barneby Award, 2004), and Sigma Xi-George Washington 

University chapter. This is contribution number 175 in the 

Smithsonian’s Biological Diversity of the Guiana Shield Program publication series.

1. P. bicolor

1a. P. bicolor var. bicolor

1b. P. bicolor var. velutina

2. P. brasiliensis

3. P. coccinea

4. P. duckei

5. P. durissima

6. P. emarginata

7. P. fanshawei

8. P. grahamiae

9. P. guianensis

10. P. induta

10a. P. induta subsp. induta

10b. P. induta subsp. glabra

11. P. leiogyne

12. P. macrostachya

13. P. paraensis

14. P. princeps

15. P. riparia

16. P. sandwithii

17. P. speciosa

Appendix A: 

Numerical List of Taxa

Appendix B: 

List of Exsiccatae

The numbers within parentheses refer to the corresponding species number in the text 

and in the Numerical List of Taxa.

Acevedo-Rodriguez, P. 3266 (3), 6044 (14), 14648 (4)

Appun, C. F. 263 (15), 857 (3), 1702 (15)

Archer, W. A. 7626 (4)

Aublet, J. B. F. C. s.n. (9)

Aymard, G. 8029 (14)

Aymard, G. & Delgado, L. 7887 (14)

Beck, H. T. et al. 139 (5)

Berry, P. E. 4797 (14)

Berry, P. E. et al. 6603 (12)

Black, G. A. 2654 (12)

Boom, B. M. & Mori, S. 1932 (14), 1940 (14), 1950 (14), 1953 (14), 1958 (14),

1960 (14)

Boom, B. M. & Weitzman, A. L. 5718 (14)

Bordenave, B. N. 871 (9)

Bordenave, B. N. & Raes, N. 5135 (15)

Cid Ferreira, C. A. et al. 916 (6), 6795 (10b)

Clarke, D. C. 168 (3), 753 (5), 1327 (3), 2314 (3), 3010 (10a), 3207 (10a), 3289 (10a),

3408 (3), 3916 (3), 4202 (14), 5068 (3), 7687 (10a), 7790 (10a), 11061 (15), 11094 

(15), 11338 (15), 11374 (15)

Cowan, R. S. 38737 (15), 38814 (15)

Cremers, G. 1328 (2), 5063 (2), 7323 (14), 9864 (9)

Croizat, L. 766 (14), 925 (14)

Davis, D. H. 815 (3)

de Lima, H. C. 2156A (13), 3298 (11)

Ducke, A. 48 (17), 113 (1b), 157 (11), 302 (14), 328 (14), 591 (13), 767 (4), 10882 (2),

10883 (2), 10984 (1b), 12245 (4), 16416 (2), 16449 (13), 16751 (1b), 20311 (4),

23285 (11), 23287 (1b), 23723 (13), 23726 (1a), 23727 (1b), 23729 (5), 23730 

(17), 29025 (14), 34959 (1b), 34960 (1b), 34961 (1a), 34962 (14), s.n., RB 10882 

(2), s.n., RB 108837 (2)
4 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Ek, R. C. et al. 614 (15), 1579 (9)

Evans, R. E. et al. 1992 (15), 1998 (3), 2327 (15), 2446 (15),

3516 (15)

Ewel, J. 104 (14)

Fanshawe, D. B. 1044 (7), 1498 (7), 4881 (15), 4912 (15)

Feuillet, C. 10065 (9)

Fleury, M. & Pignal, M. 1691 (2)

Fróes, R. L. 22282 (12), 25762 (9), 28100 (4)

Gillespie, L. J. 1579 (3)

Gonggryp, J. W. 3737 (9)

Graham, V. 197 (16), 355 (8)

Granville, J. J. de & Crozier F. 17222 (9)

Grenand, P. 633 (15), 649 (13), 1285 (15), 1781 (9)

Guppy, N. 513 (6), 172 (3)

Hammel, B. et al. 21242 (15), 21584 (14)

Henderson, A. & de Lima, J. R. do N. 604 (3)

Henkel, T. W. 1969 (3), 3566 (3), 3662 (3), 5108 (10a)

Hoff, M. 7211 (9), 7323 (14)

Hoffman, B. 979 (3), 1069 (7), 1317 (3), 1814 (7), 1897 (7),

4877 (14), 5924 (6)

Hoffman, B. et al. 1454 (15), 4759 (15), 6635 (14), 6794 (14)

Hopkins, M. J. G. et al. 741 (3)

Irwin, H. S. & Westra, L. Y. Th. 47476 (9)

Irwin, H. S. et al. 55072 (14)

Jansen-Jacobs, M. J. et al. 2273 (3), 2890 (3), 3731 (3)

Jenman, G. S. 893 (16)

Kappler 81 (9)

Krukoff, B. A. 1167 (13), 1184 (13)

Lang, H. 413 (15)

Lang, H. et al. 292 (15), 332 (15)

Lanjouw, J. 732 (15)

Lanjouw, J. & Lindeman, J. C. 2034 (9), 2851 (14)

Lewis, G. P. 1673 (3)

Liesner, R. L. 17125 (11), 18701 (14), 7128 (12), 7200 (12)

Lindeman, J. C. & de Roon, A. C. 874 (9)

Lindeman, J. C. et al. 172 (3)

Maguire, B. 40752 (13)

Maguire, B. et al. 36705 (11), 36736 (14), 36796 (14), 41952 

(11), 41986 (11), 42602 (11), 47045 (9)

Marcano-Berti, L. & Salcedo, P. 67–979 (12)

Matos, F. et al. 190 (10b)

McDowell, T. 2305 (3), 2374 (3), 3758 (15), 3895 (7), 5744 (8)

Mélinon, M. 18 (9), s.n. (9)

Mennega, A. M. V. 548 (15)

Milliken, W. & Bowles, S. 290 (14), 389 (3)

Mori, S. 24362 (3)

Mori, S. et al. 23777 (14), 25433 (14), 27358 (9)

Mutchnick, P. 520 (15), 563 (3), 672 (3), 758 (3), 987 (3)

Myers, J. G. 3906 (3), 5801 (10a)

Nascimento, O. C. et al. 148 (11), 198 (11)

Nee, M. 30573 (14), 30831 (14)

Oldeman, R. A. A. 1734 (2), 1835 (9), 2054 (9), 3030 (2)

Oldeman, R. A. A. & Sastre, C. 77 (2), 115 (9), 163 (2)

Pennington, R. T. & Johnson, I. RTP431 (14)

Pennington, R. T. et al. 400 (5)

Pires, J. M. 48561 (9)

Pires, J. M. & Silva, N. T. 1954 (1b), 1984 (1b)

Pires, J. M. et al. 51310 (13)

Poncy, O. 183 (9), 1666 (9)

Prance, G. T. & Ramos, J. F. 24350 (1b)

Prance, G. T. et al. 9287 (3), 10517 (14), 10639 (14),

10936 (14)

Prévost, M. F. 1717 (9)

Pulle, A. 154 (15)

Redden, K. M. 1080 (14), 1094 (3), 1104 (14), 1117 (3), 1161a 

(15), 1171 (5), 1178 (5), 1179 (3), 1180 (3), 1181 (3), 1722 

(7), 1733 (7), 3018 (3), 3019 (3), 3031 (3), 3038 (3), 3045 

(3), 3056 (3), 3057 (3), 3082 (3), 3088 (3), 3089 (3), 3090 

(8), 3092 (10a), 3099 (10a), 3103 (10a), 3104 (8), 3106 

(3), 3154 (10a), 3155 (10a), 3160 (10a), 3169 (10a), 3189 

(10a), 3208 (15), 3229 (15), 3233 (15), 3263 (15), 3281 

(15), 3297 (3), 3305 (15), 3307 (8), 3312 (3), 3314 (15),

3536 (14), 3714 (12), 4022 (16), 4648 (7), 5081 (3), 5429 

(7), 5968 (9)

Renner, S. S. 59 (11)

Ribeiro, B. G. S. 15.286 (713) (11)

Roberts, S. 2 (9)

Rodrigues, W. A. 8730 (1b)

Rodrigues, W. A. et al. 8461 (1b)

Rombouts, H. E. 674 (15)

Sabatier, D. & Prévost, M. F. 4674 (2)

Sastre, C. et al. 6470 (9)

Schomburgk, R. 251 (3), s.n. (3), s.n. (14)

Schultes, R. E. 24578 (11)

Sigueira, R. 34958 (13)

Silva, N. T. & Brazão, U. 60669 (14)

Silva, N. T. & Rosario, C. 3967 (2)

Silva, N. T. et al. 152 (9)

Smith, A. C. 2521 (3), 2652 (10a), 3031 (3)

Snethlage, E., s.n., H.A.M.P. 10.117 (2)

Sothers, C. A. et al. 672 (17)

Souza, R. s.n. (14)

Spruce, R. 2671 (12)

Stahel & Gonggryp 2861 (3)

Stergios, B. & Velazco, J. 14003 (14), 14256 (14)
NUMBER 109 • 4 1

Sterigos, B. & Yanez, M. 15203 (14)

Steyermark, J. A. 104038 (11)

Steyermark, J. A. & Bunting, G. S. 102923 (12)

Teunissen, M. & Schulz, J. P. 13650 (15)

Tiwari, S. 488 (10a), 511 (10a), 517 (10a)

Thomas, W. W. & Plowman, T. 3151 (14)

Thomas, W. W. & Samuels, G. 3290 (14)

Ule, E. 8146 (3)

Versteeg 248 (10a)

Veth, B. & Manu 32 (9)

von Rohr, s.n. (9)

Williams, L. 14790 (14)

Wilson-Browne, S. J. WB136 (3)

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notice des plantes de l’Isle-de-France. Didot, Londres, Paris.

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Mr. Schomburgk in British Guiana. Journal of Botany (Hooker), 2:127–146.

Bruneau, A., F. J. Breteler, J. J. Wieringa, G. Y. F. Gervais, and F. Forest. 2000. Phylogenetic Relationships in Tribes 

Macrolobieae and Detarieae Inferred from Chloroplast trnL Intron Sequences. In Advances in Legume Systematics, part 9, ed. P. S. Herendeen and A. Bruneau, pp. 121–149. Royal Botanic Gardens, Kew, UK.

Cowan, R. S. 1957. Studies in Tropical American Leguminosae III. Brittonia, 8:251–253.

Cowan, R. S. 1976. A Taxonomic Revision of Elizabetha (Leguminosae-Caesalpinioideae). II. Proceedings van de 

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